Accepted Article archiving. article maybe used fornon-commercialpurposes inaccordancewithWileyTerms andConditionsforself- 103: 1069-1098, whichhasbeenpublishedin finalformathttps://doi.org/10.1111/1365-2745.12424. This M. andFollak, S.(2015),BiologicalFloraof theBritishIsles:Ambrosiaartemisiifolia. JournalofEcology , H. ,Petitpierre,B.Richter,R. ,Schaffner,U.Smith,M.Starfinger,Vautard,R. , Vogl,G.Lippe, Kazinczi, G.,Kueffer,C.Laitung, B.,Lavoie,C.LeitnerM.Mang,T.Moser,D. Müller‐Schärer, O. ,Bullock,J.M.,Chapman, D.S.,Chauvel,B.,Dullinger,S.Fumanal,Guisan,A. Karrer, G., This isthepeerreviewedversion ofthefollowingarticle:Essl,F.,Biró,K.Brandes,D. Broennimann, Vienna, Austria; University ofNaturalResourcesandLifeSciencesVienna , GregorMendelStreet33,1180 Dynamics, UniversityofLausanne,1015 Switzerland; 11 Université Blaise-Pascal,UMR547PIAF,BP10448,63000 Clermont-Ferrand,France; Conservation &Analyses,Giessergasse6/7,1090Vi enna, Austria; 8 UK; 6 5 Institute forPlantBiology,Mendelsohnstraße4,38106BraunschweigGermany; Protection, 8361Keszthely,DeákF.str.16,Hungary; Matieland 7602,SouthAfrica; Invasion Biology,Departmentofotany&Zoology,StellenboschUniversity,PrivateBag91, Landscape Ecology,UniversityVienna,Rennweg14,1030Vienna,Austria; 1 Follak Franz Essl Franz Biological Flora theof British Isles: List Vasc.Pl. Br. Isles (1992) no. BIOLOGICALFLORA OFTHE BRITISHISLES* Article Type:Biological Flora Uwe Starfinger Uwe Müller-Schärer Laitung G Antoine Bullock INRA, UMR1347Agroécologie,21000Dijon,France; NERC CentreforEcology&Hydrology,BensonLane,Wallingford,OxfordshireOX108BB, Department ofEcology&Evolution,UniversityLausanne,1015Switzerland; Department ofBotanyandBiodiversityResearch,DivisionConservation,Vegetation INRA, UMR547PIAF,63100Clermont-Ferrand,France; 7 NERC CentreforEcology&Hydrology,BushEstate,EdinburghEH260QBUK; 26 6 16

, D , ClaudeLavo uisan 1,2,† S.Chapman aniel , Krisztin 23 19 5,12 14 , Robert , Blaise Petitpierre , Blaise Kaposvár University,FacultyofAgriculturalandEnv ironmental Sciences, , G Karrer erhard a Biró ie Vautard 17 , Michael Leitner 3 7 3 University ofPannonia,GeorgikonFaculty,InstitutforPlant , Bruno Chauvel , D 135, 74, 1 ietma 24 , Gero Vogl 5 13 , Robert Richter , G r Brandes

abrie 18 Mang , Thomas la Kazinczi 8 , S 20 4 , Moritz vond , Dullinger tefan O 4 Technical UniversityBraunschweig, livier Broennimann livier 9 20 Vienna InstituteforNature , U 12 14 rs Schaffner rs Institute ofEarthSurface , Chri 1 13 , Dietmar Moser , Dietmar L er Institute ofBotany, 10 stoph Kueffer 1,9 Clermont Université, , Bori ippe 21 , Matt S 2 25 5 s Fumanal Centre for , James M. andSwen No. 278 15 1 mith , Heinz , Beryl

10,11 22 , , Accepted Article and management. management. and phenology, floral and seed characters, herbivores communities, responses to biotic factors, responses to environment, structure and physiology, within thestandard frameworkofthe (Common ragweed)thatare relevantunderstanding to ecology. topics its main The are presented Vienna, Austria; Poland; Universitätsstrasse 16, 8092Zürich, Switzerland; Fribourg, Switzerland, Biology, Unit Ecology & University Evolution, of Fribourg, Chemin du Musée 10, 1700 1. Summary Europaea of Nomenclature vascular plantsspecies, non-British for Staceand, follows (2010) * [email protected]: †Correspondence author. Running head:Ambrosia artemisiifolia Austria Safety, Institute for Sustainable Plant Pr Québec City, Québec, G1V 0A6 Canada; supérieure d’aménagement du territoire et de développement régional, Université Laval, 22 Universität, München, Lichtenbergstraße 1, 85747 Garching, Germany; This isarticle by protected copyright.reserved. Allrights 24 40, 7400 Kaposvár, Hungary; ofPlantScience,Institute Department ofPlantProduction andPlantProtection, Guba S. str. Agroécologie, AgroSup/INRA/uB, 17Agroécologie, rue AgroSup/INRA/uB, Sully,21065 Dijon Cedex, France; National National and International Health,Plant Messeweg 11/12, 38104 Braunschweig, Germany; 12165 Berlin, Germany; and Germany; 12,12165Rothenburgstrasse Berlin, Gif sur Yvette Cedex, France; Laboratoire des Sciences du Climat et de des Sciences de Laboratoire duClimat et Laboratory of Aeropalynology, Faculty of Biology, Adam Mickiewicz University, Pozna This account presents information on all aspectsthe biology of artemisiifolia Ambrosia of 23 . Julius Kuehn Institute, Research Institute Federal for Kuehn forJulius Cultivated Centre Plants, 21 CABI Switzerland,CABI Chemin des Grillons 1, 2800 Delémont, Switzerland 20 Faculty ofPhysics,University Vienna, Boltzmanngasse5,1090 15 Institute of Integrative Biology, ETH Zürich, of ETHZürich, Biology, Institute Integrative 25 Institute Technical of Ecology, University ofBerlin, Biological Flora of the British Isles

18 Heinz Maier-Leibnitz-Zentrum, Technische l’Environnement, IPSL,CEA/CNRS/UVSQ, 91191 oduction, Spargelfeldstraße 191, 1220 Vienna, and disease,and history, andconservation, impacts 16 Université deBourgogne,1347 UMR 26 Austrian Agency for Health and Food Austrian Agency forHealthand 19 : distribution, habitat, habitat, distribution, : Department of 17 École École Flora L. ń , Accepted Article This isarticle by protected copyright.reserved. Allrights biology geographical and altitudinal distribution, health, modelling, parasites and diseases, reproductive Key-words: Europe. consensus among models that climate allowitsnorthwardchange and will spread in up-hill its copious, highly allergenic pollen isThere public healthproblems. creates considerable 7. reducesubstantially growth andseed production of southern This Switzerland and northern Italy. species capacitytohaveappearsto the 6. Europe and is now inabundantopen, habitats disturbed as anda ruderal weed. agricultural widespread regions intemperate become II,it has of War century.in the19th World Since 5. rich nutrient level. It ha the growing season. terminate autumn frosts 4. tofrom summer autumnOctober). earlyto (July Intemperate thermalrange. wide c European spring, rateof the vegetativeon growth temperature, depends butdevelopment occurs over a 3. pistillate florets (female) are in arranged groupsaxils theof andlateral main stem leaves. composed of staminate (male) florets terminatestems, the while cyme-like clusters of has erect, branched stemsandpinnately leaves. lobed racemes Spike-like of malecapitula whoseAmerica height varies from 10 to2.5cm m accordingconditions. toenvironmental It 2. Ragweed. , tribe Heliantheae. leaves mostly opposite, alternate deltateleaves mostly above, to opposite, to 2.5 stemsm tall; hairy, bluntly erect, 4-angled, ridgedleafy, and with branches; several monoecious herb,annual which the germinates in spring andsets fruit in the autumn. Plants Ambrosia artemisiifolia artemisiifolia Ambrosia

In heavily infested regions of Europe, Recently, the N. American ragweed leaf beetle ( Recently, the N. American ragweed leaf beetle Seeds requireSeeds prolongedbreak chilling to Ambrosia artemisiifolia “ Ambrosia artemisiifolia agriculture, biogeography,ecophysiology, change,climate agriculture, germination, [“ Ambrosia artemisiifolia was introduced intoEurope with seed from imports North America is sensitive to freezing. Late spring frosts kill kill seedlings and first spring frosts the to freezing. Late sensitive is is a monoecious, wind-pollinated, annual herb native to North North to is a monoecious, wind-pollinated,native annual herb wird sich in Mitteleuropa wohl nie in gefahrdrohender Weise vermehren“ vermehren“ Weise gefahrdrohender in nie wohl Mitteleuropa in sich wird A. artemisiifolia will never will inbecome dangerouslyCentral abundant Europe“] Ambrosia artemisiifolia artemisiifolia Ambrosia limates, and female male flowers areproduced dormancy. Following seedling emergence in s a preferences a dryintermediate for of to soils lanceolata or elliptic in outline, 25-55(90) x 25-55(90) outline, in elliptic or lanceolata Ophraella communa A. artemisiifolia causes substantial crop-yield losses substantial losses crop-yield and causes

. L.is awindpollinated, ) hasbeen detected in (P. Ascherson 1874) Accepted Article increasingly apparent in southern parts of the British Isles. in healthpublic Centraconsiderable problems 2003; Sheppard oncropyields (Reinhardt impact toits economic Inaddition agricultural ofcrops. weed habitats; inopen, ruderal native bothits co and This isarticle by protected copyright.reserved. Allrights projections. Male flowers produce copi spiny in ending ridges (4)5-7(8) longitudinal with syconium ahairy forming involucrum reddish-brown,± to each to its yellowish adnateovoid, achene Fruit in woody achenes. enclosed x1.8-2.5 mm,3-4 mm c. with single flower.Seeds spines mm) (eachlike each 3-5 bristle- with5-8 small cupule-like byinvolucre bracts; groups, surrounded small in2-5 erect Fema 12-20,anthers free. florets finely hairy, mm) male. Male flowers with cream, ye spike-like ebracteate racemes, femalein capitula axillaryimmediately below the clusters stalkedshort indense, male two kinds: elongatingcapitula of Inflorescences gland-dotted. sparsely finelywith stiff hairy slender bris or deeply20-30(50)bases pinnatifid, mm, cuneate,or margins toothed; entire facesabaxial tenuifolia, A. A. trifida tenuifolia, regionand tropical Africa),whereasothers nativetoNorthAmerica three ( 2006). OnlyCommittee Editorial centre of diversity comprises the south-western USA and adjacent northern Mexico (FNA species ( species populations haveEuropean beendistinguished artemisiifolia A. from range and abundance since the mid-20th greatly where ithas Isles and continental includingin increased widely, Europe, theBritish artemisiifolia Gaudeul c The genus In the British Isles, apitula; involucres campanulate or turbinat involucres or campanulate apitula; A. elatior et al. et is restricted is North America, to has it colonized temperateregions world ofthe 2011) conspecific their confirm Although status. range native the of et al.et containsAmbrosia species, leastat 40 numerous intraspecificwith taxa.Its L.; see Tropicos 2014).However, genetic studies (Genton ) are established in Europe (Rich 1994; Fumanal 2007; Follak 2007; 1994; Fumanal (Rich Europe ) are established in 2006), the strongly allergenic pollen of thestrongly common pollen allergenic ragweed 2006), causes Ambrosia artemisiifolia L. is native to the Old World (Mediterranean the (Mediterranean L. Old to native World is maritima A. llowish or pale green corolla in drooping short-stalked (2-5drooping short-stalked in pale llowish orcorolla green ous amounts ofwind-dispersed pollen.

century. tles (strigillose), adaxial faces strigillose, both le flowers without corolla or pappus, capitula corollaorpappus, le without flowers l andEast Europe, andthese becomingare ntinental European itis European ranges an important ntinental e, 2-3 mm in diameter, glabrous, indiameter, hispid 2-3mm e, is primarily an alien invasive plant of plant invasive an alien primarily is

s. str. s. A. psilostachya, A. A. as a distinct distinct a as et al. et et al. 2013). 2005; et al.et or A. A.

Accepted Article Russia (CABI 2014). In Russia, it had invaded an area of more than 50,000In (CABI invaded km more than had 2014). Russia, areaofan Russia it 2008a). Further east, populations largest in the PannonianPlains of most species widespreadis southern ofwith parts in Eastern 2), Central the Europe and (Fig. the 20 This isarticle by protected copyright.reserved. Allrights Brandes & Nitzsche 2007;Tokarska-Gudzik Chauvel Prots 1998; & Song century (e.g. mid-20th the status ofthe species in Central andSouth America. are theredetails andinvasionpossibly but onthedistribution few Uruguay, Paraguay and 2014), Nakajima and & (Mondin Brazil Strong (Acevedo-Rodríguez 2007), Hispaniola & 2014), Chile(established; Fuentes N. unpublished data), the Bahamas andthe island of speciesto Argentina the spread (Anton&Zuloaga has IntheAmericas, 1987; EPPO2013). (Lazarides Australia data) unpublishedHenderson Watanabe Japan (widespread; & EPPOwidespread; Duan NakayamaChen 2000; 1998; 2013), mid-19ththe Since century, especially decades, and recent in it is (where China including Europe, world regions the of temperate several invaded in southern England, in particularin urban areas (e.g. London). whereas in itisrare Ireland (L In Britain, distribution andaltitudinal Geographical I. and in thenorth of far Europe. Bullock populations smallare and ephemeral.often Itis distributed in the northern regions of Cent (Chauvel (Moskalenko 2001). repeated colonisations(Chapman isScandinavia, unclear but it to what extent th In Europe, In Europe, cent and southern hotspots are invasion Other century and it was still spreading at the very beginning of the 21 the beginning very of spreading the at wasstill it century and et al. et al. Ambrosia artemisiifoliaAmbrosia 2006), and the Po plains of northern Italy. 2002; EPPO 2013), South Korea (EPPO 2013), South Africa (established; L. (established; SouthAfrica (EPPOKorea 2013), South 2002;EPPO 2013), Ambrosia artemisiifolia A. artemisiifolia A. ockton & Crocker Fig.1). records Most ockton 2014; been have made et al. hasis scattered occurrencesexpanding but distribution, its is also widespread in Ukraine (Song & Prots 1998) and Prots 1998) (SongUkraine & widespread is in also 2014). has greatly increased in range and abundance since since abundance and range in increased greatly has these these naturalisedor transient populationsare and et al. et al. et

ral, Western and Eastern Europe, andmost EasternEurope, ral,Western and Hungary, Croatia, Croatia, Hungary, and Serbia (Kazinczi largely absent from Mediterranean Europe Mediterranean absent largelyfrom (2012)collatedrecords as farnorth as 2011; Bullock ral France, theRhône in particular valley et al.et

1997),Zealand (Webband New Ambrosia artemisiifolia et al. 2006; KissBeres 2006; 2006; & et al. 2012). Currently,the A. artemisiifolia 2 by the byend of the is patchily st century etal. has

Accepted Article II. Habitat II. Kleinbauer 2009; Cunze 2009; Kleinbauer Dullinger growing & that are sumsduring characterized by period(Essl, the high heat Similarly, inEurope, the species hasitsoptimum& Crompton intemperate climates1975). range, native In its (A) CLIMATIC AND TOPOGRAPHICAL LIMITATIONS This isarticle by protected copyright.reserved. Allrights stratification (Shrestha (Shrestha stratification constrains growth (Chapman Mediterranean climate possibly because low severly rainfall months during summer the Europe, In Europe. southern parts of higherelevations in most from of commonragweed explain also the absence requirements 2009),Kleinbauer as the species not is able to complete its life cycle. These thermal & Dullinger (Essl, factor climatic important limiting the as identified most been has months 2007;de Merlis Simard & Benoit2010; Joly GoursaudLavoie, Jodoin & 1975; & Crompton (Bassett soybean fields maize and crop and in and widespread in southern Ontario andQuébec, especially along railways andpaved roads, species the abundance Canada, of distribution but in spatial and (Brouillet Territories New Brunswick, Nova Scotia, Prince Edward Newfoundland,Island, and in the Northwest greatlyexpanded 20th during the be itinunlikely Columbia,century, native to and British is Nevertheless, Goursaud2007). deMerlis Jodoin & (Lavoie, thought previously extensive than native wasmore the range that suggests Québec, southern been which sincehave from specimensrecovered Older specimens. wasbase 1975),butthisassertion & Crompton that been argued has It 2006). Committee Nunavut Editorial (FNA the Yukonand of exception with territories, reconstruct. In Canada theand USA, the species ispresent in all provinces, states and to difficult range ofragweed’s is common native the extent settlers, bycolonization Canada. range North However,following within expansion to historical America due

Ambrosia artemisiifoliaAmbrosia

A. artemisiifolia A. artemisiifolia A. et al. et al. et al. 2014). Except for Québec,2014). for Except recentthere is no documentation of the 1999). et al. et 2013). In Central Europe, low 2013).In m Europe, Central nativeis and widespread inthe United States and southern is native only to the central part of North America (Bassett NorthAmerica of part is nativethe only central to 2014), winter temperatures too and allow seed highare to growsbest under temperate, climates continental (Bassett A. artemisiifoliaA. et al. common ragweed’s North American range American common ragweed’s North 2011;Ngom& Gosselin 2014). d on a few old(19th few d ona herbarium century) is largely absentfrom regions a with

ean temperature summer ofthe A. artemisiifolia isabundant Accepted Article Polygonum and of trampled vegetationPlantaginetea, ofclass (e.g. Polygono-Poetea (i.e.summer annuals suchas European range(Table 1) speciesare diagnostic of segetal vegetation ofthe class Stellarietea most commonly associatedspeciesrecorded Easternits in relevés within and Central Although Communities III. 2010). (Landolt(BOKU Similarvaluesfor Switzerland 2014). given are fertility) high soil undermoderately cond best basic pH(grows soil for 8 dry for soils), preference (indicating a moisture given3 for are of values This isarticle by protected copyright.reserved. Allrights communities ofnatural habitats (e.g. prairiegrassland) (Bullock PlantBritishCommunityin the classification ofRodwell (2000). S1; Brandes NitzscheEurope (Fig. 2007; & Bullock et al. soils inNorth America (Bassett & Crompton 1975). in Europe(Fumanal Ambrosia artemisiifolia SUBSTRATUM (B) Nichols 2012) Nichols urban ruderal and (Ziskahabitats inarable WebsterfieldsKotanen (e.g. 2007;MacKay & 2008), Merlis Goursaud& de in other such habitats gravelas bedsof ri and construction sites, embankments, railway roadsides, e.g. habitats to ruderal restricted itislargely Isles, the British such as invaded, less In currently regions 2014a). Fiedler& Karrer Milakovic, arable fields andonroadsides (Essl, &KleinbauerDullinger 2009; Pinke communities of disturbed habitats communities of Nature Conservation 1999) or for Central Europe (Ellenberg 1999) or Europe Central for In accordance with common ragweed’s habitat preferences, largea fractionof the No Ellenberg indicator values for valuesfor No Ellenberg indicator In the more heavily infested areasof Europe,In heavily more infested the Ambrosia artemisiifolia Ambrosia

aviculare

et al. agg.). In addition, diagnostic species ruderal of vegetation dominated by ). In its native range, range, native Inits ). landfill sites sites (Bullock landfill colonizes ofwithout awiderange preference soil types anyparticular 2008), although seems it to show a preference for clays orsandy Chenopodium album, Conyza canadensis colonizes wide rangeofa disturbed habitats Britain in and itions), and 6 for nitrogen level (indicating intermediate to intermediate (indicating and 6fornitrogenitions), level - roadsides,along inwastelands Jodoin & (Lavoie, A. artemisiifolia vers, semi-dryor disturbed grasslands (see et al. et A. artemisiifolia A. et al. et 2012). The species is occasionally recorded 1992). 1992). However, for Austria, Ellenberg et al. et et al.et A. artemisiifolia artemisiifolia A.

are available for Great Britain (Hill Britain Great for available are 2003) 2012), it was innot recorded any frequently tothe contributes et al. - but rarelyis found in 2012). and is most frequent in Plantago major,Plantago Setaria Setaria pumila et al. 2013; XI. )

Accepted Article This isarticle by protected copyright.reserved. Allrights perennial species (class Artemisietea; e.g. heavily invaded regions (Bullock (Bullock regions invaded heavily al. highly competitive and can cause significant yi fields, such arable as incontinuously habitats, 2007). However, disturbed dist after occurs alongrivers) usually only et al. (B closesvegetation duringsuccession as the As anannualofopen habitats, IV. Response to biotic factors artemisiifolia lanceolata Plantago repens millions of individuals), some at (up 400 m ofextremelydensities individuals), at highto plants some millions most populationsin less sma areas invaded are Ambrosia artemisiifolia (A) GREGARIOUSNESS V. Responseto environment (See sites invasiveness its may inmoist conditions. field success limit competition Interspecific under Gard often wilt above the fracture (Brandes & Ni Bullock easily(Nitzsche to break 2010; individuals mature tend and growth of inhibit germination than rather donorplants-promote leaves sunflowerfrom -and other of (see rare are species in shootseven after 90%ofthe leafarea had been removed. artificial which defoliation, did not repr affect 2008b; Novak 2008). Colonization of closed vegetation types (e.g. semi-dry grasslands, embankmentstypessemi-dry grasslands, (e.g.closed vegetation Colonization of 2008). et al. ) and of fertilized grasslands (class Molinio-Arrhenatheretea; e.g. e.g. Molinio-Arrhenatheretea; of (class grasslands fertilized ) and Ambrosia artemisiifolia Ambrosia

Reports on the susceptibility of of susceptibility onthe Reports V. (2013) showed that both native andintroduced bothnative that invasive (2013)showed C). . A. artemisiifolia A. et al. VI. VI. and 2009; Bullock populations populations vary insize greatly andspatial extent Whereasin Europe. Trifolium repens Trifolium F). Incontrast, Kazinczi

shows little tolerance to trampling, because the main stemsbecause main of the to tolerance trampling, little shows Ambrosia artemisiifolia , and they speculate that this may toits contribute invasion et al. et al. 2012). A. artemisiifolia A. 2012). ) are found frequently in relevés containing containing inrelevés frequently found ) are Artemisia Daucus vulgaris, carota urbance hasNitzsche createdurbance & gaps (Brandes azzaz 1979; Brandes & Nitzsche 2007; Fumanal 2007; Nitzsche azzazBrandes & 1979; oduction, oduction, and plants reallocatecould resources tzsche 2006). Under experimental conditions, Underexperimental 2006). tzsche eld losses,row especiallycrops in (Kazinczi ll and ephemeral,ll veryand et al. is by plant replaced perennial species (2008d) found that that (2008d)found aqueous extracts to allelopathic effects of other plant ofother effects to allelopathic

et al. 2012); thebroken stems A. artemisiifolia artemisiifolia A. large populations(with large Medicago lupulina, Medicago A. artemisiifolia A. -2 ), prevail in and Elymus tolerate tolerate A. A. et is Accepted Article This isarticle by protected copyright.reserved. Allrights mass, i.e. on average, seeds are larger in Europe. the native range.Differences in frost tolerance attributed were partly to differences in seed seedlingsthat of common ragweedhave higher European frost in tolerance the rangethan in Fumanal 2004;La Crompton1975; DiTommaso (Bassett & inurban wastelands, particularly and cultivated sunflowerfields, among maize, andsoybean and invasive along ranges: waterways, roadsides artemisiifolia Ambrosia HABITATS VARIOUS IN PERFORMANCE (B) the growing season(Ziska autumn terminate frosts the first 2014) and Kaviani &Tackenberg seedlings (Leiblein-Wild, artemisiifolia Ambrosia (C) EFFECT OF FROST, DROUGHT, ETC. been found in Québec2004;Eom,& (DiTommaso DiTommaso Weston2013). roadside plant species (Joly of salt tolerant selection ecotypes of tode-icing contributed have saltmay ofquantities largethe of in application Inaddition, roadsides Québec. have of the colonization facilitated networkduringthethe improvement (paving) 20th orthe theroad extension of century may Jodoin& 2009).Lavoie, Kleinbauer & Dullinger only the1960s)colonized (since (since roadsides 1970s) recently: fields (Essl, the and places, etc.) were invaded. Those habitats curre contaminated crop seeds. Subsequently,ruderal habitats distantrailways to feeding (bird patternhas 3):railw beendifferent observed (Fig. increasingly railways (ChauvelroadsFrance and along in 1920s(Kazinczithe asearlyin weed of occurrences first Similarly, America. North mid-20th theuntil asitwas century, primarily Goursaud deMerlis 2007). In France, Jodoin1930s)theroads railways 1920s), (Lavoie, the (since and & (since fields invaded expansion its range of In habitats. Québec, it spreadfirst along river corridors, and later During its spread inEurope and During ragweed NorthAmerica, has spread common its undergone an et al. 2008; Simard & Benoit2010;Bullock thrives in a wide range of open and disturbed habitats, in bothnative habitats, in anddisturbed open of range thrivesawide in plants are sensitive to freezing. In particular, late spring frosts kill spring frosts late particular, In freezing. to sensitive are plants et al. et al. 2011). Leiblein-Wild, found2011). Kaviani &Tackenberg (2014) 2011). However, to date, to 2011).However, salt tolerant ecotypes have only A. artemisiifolia A. artemisiifolia et al. 2008a). However, 2008a). from the 1950s, itwas found A. artemisiifolia A. introduced inintroduced contaminated imports seedfrom ntly with the ntly havelargest withthe populations been and railways, in old or in and railways, fields andindustrial this expansion. This might have led to the might This expansion. led have this ays of wereinvadedfirst, throughspillage Goursaud de Merlis (2007) suggested that (2007) suggested Goursaud deMerlis voie, Jodoin & Goursaud de Merlis 2007; &Goursaud deMerlis Jodoin voie, with a with competitive advantage over other et al. wasmostly confined to crop fields

et al. 2012;Ngom & Gosselin 2014). 2006). In Austria, a somewhat a 2006).In Austria, in Hungary asan were arable Accepted Article (A) MORPHOLOGY (A) MORPHOLOGY VI. Structure and physiology the invasionlimiting of sites. moist &(Leiblein Lösch which 2011), suggests that competition main interspecific is factor a photosynthesis rates under mois conditions. field under waterlogged sites occur in waterloggedand tolerates contenthigh water roots and18% to stems (Nitzsche 2010). by 2010). drySide-branches(Nitzsche followed 27%to54% of representallocation, the mass (Karrer damage Additional shoots candevelop accessoryfrom buds, which areusually induced by physical numerous node)andto thethird includes side-b the 2–4cmabovesoilfirst level(at internodes. about at Branching the by starts of elongation Plants continue and4 cm. between diameter 0.3 appressed and/or shorter, of Number hairs. todenselysparsely with relatively pubescent withrepr positively correlated vol suchheighttraits asand life-history 0.1 varies from to 2.5m toenvironmental conditions and according Vegetativecompetition. and shows (Fig. 4), parts height only Its apical &Aarssen1996). (Irwin aweak dominance artemisiifolia Ambrosia This isarticle by protected copyright.reserved. Allrights and seedling establishment (Shrestha on impact high negative disproportionally springhasa stress in quantities(Raynalsmall Leiblein &Bazzaz &Lösch 1975; 2011).Furthermore, drought in albeit seeds, produce able to remain but growth, stunted have plants ragweed common damage (Almádi irreversible without content in thatwater-saturated common ragweed leaves lose upto70%can of their maximum water (Bazzaz 1973; Bazzaz 1974). the plants 1973), but content (Bazzaz soil water The net photosynthetic of rate The net Although under experimental conditions conditions experimental Althoughunder et al . 2011). The stem breaks easily b is an annual erect plant, which is usually highly branched in the upper in the usually branched highly is which plant, erect annual is an Ambrosia artemisiifolia

oductive traits(e.g. seedand t soil conditions in the ofabsence plant other species et al. A. artemisiifoliaA. 1999). ume plants,of shootnumber andbiomass are nodes ranges between 6 and 23, andbasal stemnodes 6and23, ranges between conditions (Leiblein & Lösch does & 2011), it (Leiblein notconditions long pustularbases hairsusually minute, with 1976). In or years sites,unusually dry1976). on dry to grow after the toonset ofthe but grow after flowering, only A. artemisiifolia recover rapidly from short-term droughts droughts recover rapidly from short-term Ambrosia artemsiifolia artemsiifolia Ambrosia ranches (20-29) of first to third order. third to first (20-29) of ranches tolerates a high water saturation deficit, water high saturation a tolerates decreases during reduceddecreases of periods

ecause internodes the are hollow pollen production). Stems are A. artemisiifolia A. with can germinate in soils has thehighest net germination germination Accepted Article not or slightly flattened. Seeds are spiny ridges in projections of ending 4-7longitudinal series a single develops syconiumhairy to ovoid involucrumits andadnate 6).Thesparsely asyconium (Fig. forming moderately to developswhich one-seeded to a (achene). fruit fruiting,At it is 3–5 mmlong, orlessmore lobesglabroussparsely that3–9 shallow are or is wide, with headsStaminate and thestaminate involucre 2–4mm spike-like in racemes, are the small (pistillate) are groupsfemale solitarycapitula orin upperin theaxils the of leaves. upper surface. usually has also is longer main hairs oronly not, than and paler veins, the along the slightly, hairs and sometimes appearing somewhat grey the surfaces sparselytoothed, to moderately tofew-lobed Theentire). ultimate lobes lanceolateare to narrowly entire oblong, or few- 3 pinnatelytimes lobed with more than 5 pr ovate to broadly ovate in (the uppermost outline leaves are sometimes lanceolate to linear), 2– median leaves, varying obviouslyfrom petiol tothe basal petiole from diminishes the length ofthe The petioles. narrowlywinged to long, 1991) and that that 1991) and shown that disturbed habitats (orchard, old fallow). Expe habitats, but riverbanks, showed notfields) hi withcorrelated thehabitat Populationstype. highlyfrom disturbedhabitats (roadsides, ruderal constrictum G. arbuscularmain were studied by colonized three mycorrhizalfungi (AMF): 1988; & Koide Boerner its Li French 1991). In artemisiifolia Ambrosia MYCORRHIZA (B) This isarticle by protected copyright.reserved. Allrights 2006). In turn, ragweedcommon is able to bothstimulate germination of report evidence report evidence kin of selection in juvenile mycorrhizal network in that plants grown with siblings have greater mycorrhizal network plants greater grownin siblings have that with network mycorrhizal (Schreiner &spores Koide 1993) root colonizationand (File Leaves are opposite toward the stem base, but alternate toward the stem tip, with short with tip, stem toward the base, alternate stem but Leaves the oppositetoward are The numerousThe (staminate) male capitula are arranged inleafless racemes (Fig. 5) but Glomus etinicatum , and , and G. intraradices Scutellospora is considered obligately mycorrhizal in its native range (Crowell & native (Crowell range inits obligately is mycorrhizal considered increases plant growth and developmental rates (Fumanal (Fumanal rates developmental and growth plant increases facilitates phosphorusin uptake sp. (Fumanal c . 3–4mm long, 1.8–2.5mm wide(Belcher 1985). pubescent with short, somewhat broad-based somewhat with short, pubescent imary lobes uppermost(the leaves sometimes ate tosessile. Leaf blades are 3–10cm long, in colour (Tropicos 2014). colour(Tropicos in undersurface The gher less thanthosefrom AMF colonization introduced range, 94% of populationsintroduced range, 94% 35 field A. artemisiifolia riments using riments fungalvarious have inocula et al. et hairy. Each involucre one Eachinvolucre encloses floret hairy. 2006). Fungalcolonization 2006). levels were

et al. A. artemisiifolia A. plantsthrough acommon 2012). File Glomus mosseae G. intraradicesG. (Koide & Li (Koide et al. (2012) et al. ,

Accepted Article the stomata contribute tolower assimilation rates under favourable less (e.g. dry, Lösch (Leibleinconditions & A 2011). lower openingofchlorophyll content insufficient and This isarticle by protected copyright.reserved. Allrights Chromosome number reported for (D) CHROMOSOMES wind-borneproduction of & pollen(Friedman 2011).Barrett of colonizing populations of high in rates outcrossing high. Themaintenance weretobeuniformly Canada found artemisiifolia A. et al. sizes.However, Li initial population small frequently invading have as species range, native non- in the selfing of evolution natural favour the may selection that argued It hasthusbeen may plants experience an Allee because effect 2011).of &Barrett pollen limitation (Friedman (Friedman& rates Barrett 2008).Smaller moand America In North (C) PERENNATIONAND REPRODUCTION content)nitrogen which couldresult inhigher survival fecundity.and mycorrhizal associationshort-term benef exhibited sizes thanthoseroot colonization rates and grow Payne (1976) and Martin and Martin Payne (1976) predominant,whereasare are female flowers influences photoperiod the ratioof the ofsex fl temperature (see days &Crompton Beres(Bassett 1975; 1994; Kazinczi Ambrosia artemisiifolia DATA (E) PHYSIOLOGICAL 2014).Tropicos available North America from and theintroduced rangeinEurope and Asia (seeEast individualstetraploid North American are in found populations. Chromosome counts are content (up to 8.8, SD: ±0.6SD: 8.8, mgg (up to content (2012) have demonstrated (2012)have thatnoshift from Under experimental conditions, common conditions, ragw Under experimental ’s invasion of China. Similarly, outcrossing rates of common ragweed in common of Similarly, outcrossing rates China. ’s invasionof II. A. artemisiifolia A) are the main factors controlling growth and development. The length The main controllingfactors anddevelopment. A)arethe growth is a C3 plant and typically completes its life cycle within 115 to 183 typically cycle 115to life and its C3plant within is completes a et al. A. artemisiifolia A. (2014) the diploid state is frequent, but sometimes triploid and but sometimes is diploidstate frequent, (2014) the -1 A. artemisiifolia A. is strongly self-incompatible and has high outcrossing andhasoutcrossing highstrongly is self-incompatible ), CO ), 2 uptake and photosynthetic rate under moist soil rateundermoist photosynthetic uptake and is likely to be facilitated tobefacilitated by likely is prodigious the favoured favoured as the days shorten (Allard 1943). owers: under longowers:flowers day-conditions male n with Thesesiblingsnon-kin. greater having outcrossing toselfing hasoccurredduring its (e.g. fewer root lesions, higher total leaf higher total rootlesions, fewer (e.g. its re isolated populations wind-pollinatedof is is 2 eed leaves have the highest chlorophyll

n = 36(Stace 2010). According to et al. 2008b). Photoperiod and Accepted Article 1 39% of that outside, approached photosynthetic light saturation at a PAR of c. 300 µmol m thatoutside,39% of approached photosyntheticlightatPARof a saturation 300 µmol c. shade.moderate Glasshouse-grown whichplants, hadexperiencedlight availability onlyc. °C and the high rate of photosynthesis(3 g ratewas high accompanied rateof°C byand the H a high transpiration grown, mature mature grown, field- experiment, a further (Bazzaz In photosynthesis 1973). of rates higher had five-fold allergenicity (Singerallergenicity (Ziska yieldand pollenproduction seed photosynthetic rate at optimal light flux, temperature, water potential and 300 ppm CO ppm 300 potentialand water temperature, lightflux, at optimal rate photosynthetic produce an equivalent ofamount biomass dm This isarticle by protected copyright.reserved. Allrights waterlogged) ragweed conditions. Common grows best sunlight,in full was 35 c. mgdm well as triterpenoids of the the of triterpenoids as well sesquiterpene lactones, including psilostachyn, isabelin, ambrosin, cumanin andperuvin, as A. artemisiifolia DATA (F) BIOCHEMICAL production by61% (Wayne Elevated atmospheric CO atmosphericElevated of stratum biomass theplants, thus totheof top compete light. abilityincreasing to for their 2008). Thesesquiterpene lacton activities (Brückner, Lepossa & Herpai 2003). and possess antibacterial, antifungal, antiprotozoal, andschistosomicidalanthelmintic Vidotto, Tesio & Vidotto (2013) showed under laboratory and greenhouse conditions that conditions greenhouse and laboratory under showed Vidotto(2013) Tesio& Vidotto, of plantseedsgermination testedLepossa (Brückner, Herpai 2003).recent & study,In a male inflorescences, achenes). The extract of male inflorescences had the highest on impact inthe inhibitionintensity parts plant of extracts variation among different of (roots,leaves, 2003;LehoczkyHerpai Lepossa & 1996; Brückner, Zhu 1984; & Wang (e.g.species& Quijano Fisher plant different , whereas field-grown plants did not saturate at c. 730 µmol m , whereas field-grown did saturate plants not µmol at c. 730 -2 h -1 Leskovšek These substancesThese produced by ) (Bazzaz 1974).Consequently, A. artemisiifolia A. contains various secondary metabolites et al. -2 hr et al. -1 (2012a) showed that high nitrogenlevels lead toa greaterallocation (Bazzaz 1974). Their optimal temperature temperature (Bazzaz photosynthesis for was 20optimal 1974). Their et al. 2 concentrations led to concentrations ledto ingrowtha marked increase (Bazzaz 1974), α 2005). A doubling of the CO et al. - and - and 2011;Vidotto, & show Vidotto 2013).Experiments Tesio es are characterizedan by plants did not saturate at a PAR c. 1450 µmol m plants did not aPARsaturate at c. 1450 µmol 2002). β -amyrine type -amyrine type and derivatives of caffeic acid (Soluji A. artemisiifolia A. A. artemisiifolia A. than maizethan (Bassett &Crompton 1975). et al. 2003) as well as a heightened pollen2003) as aheightened as well have an inhibitory effect on growth of have on an inhibitory effect

utilizes substantially more waterto substantially more utilizes α with antibiotic properties, suchas properties, antibiotic with , 2 β concentration stimulated pollen concentration stimulated -unsaturated -unsaturated -2 -2 s -1 ; field-grown also plants

but it growsbut it wellin γ -lactone moiety-lactone 14 -2 -2 s ć -1

et al. ; the -2 -2 2 O s 2 -

Accepted Article (Deen (Deen bei solstice, withreproductive development 2006; Ziska of Europe, parts floweringinvaded earlylate Nitzsche begins August July to & (Brandes opposite leavesappearance tothe of alternateleaves. nativeIn range, the in heavilyand 1998b), but development occurs over a wide thermal range (8 - 30°C) (Deen seedling emergence, therate of vegeta (Baskinproportion germinates & later Baskin 1977; Bassett &Crompton Following 1975). April), smallmid-springand March althoughearly in a in Europe occurs to (late Germination This isarticle by protected copyright.reserved. Allrights that kill or October plants (Dahl, Ziska the &Wihl 1999; Strandhede ofTermination 2011). & flower (Friedman Barrett femalemale and flowering forplastic, example shading is triggersflowering male earlier flowering (Deen female the first days before afew occurs usually flowering male Thefirst are present. sexes artemisiifolia Ambrosia VII. Phenology A. artemisiifolia of invasion the facilitating role in might a they play that research suggests current but tested, mixturesoil (alluviumsoil andsilica sand). of 3g residues incorporation of of (90%) after Among species, testedweed sensitive crop species as growth wasby reduced than 50% more compared tothe control. seedling growth of crop andweed species. Tomato ( of material plant dead of residues 2013). This2013). also marks the of end theperiod for developmentof viable mature seeds. found in Europe (Genton inEurope found been clines inflowering phenology latitudinal & Sweet have 1971).Similar 1943; Dickerson plants latitude demonstrated that northern from acrossis significant itsrange.there variation studyusingcommonseeds A garden native et al. The onset of reproductive development is marked by a switch from the production of the a switch from by ismarked reproductivedevelopment of onset The Although the phenology of of Although phenology the 1998a). 1998a). Flowering is withmonoecious over et al. et

. et al. 2011). Flowering is triggered by declining day lengths after thesummer lengths after by Floweringday declining 2011). triggered is 1998b; Friedman & Barrett 2011). However, the relative timing of timing therelative However, 2011). Barrett & Friedman 1998b; ’s life-cycle is typical of a short-day annual (Deena short-daytypical’s annual of is life-cycle et al. Digitaria sanguinalis Digitaria 2005; Chun Ambrosia artemisiifoliaAmbrosia A. artemisiifolia A. tive growthtive dependsontemperature (Deen ng delayed at photoperiods longer than 14 hours at longerng photoperiods than14hours delayed et al.

Allelopathic effects in have the field notbeen s flowered earlier in a northern garden (Allard (Allard garden northern a in earlier s flowered A. artemisiifolia A. 2011; Hodgins & Rieseberg 2011; Leiblein Hodgins& Rieseberg 2011; 2011; ing occurs with frosts with latefrom ing September frosts occurs suffered a large germination reduction in in soil affected the germination theand soil affected germination in Solanum lycopersicum lap in the period when flowers of both intheperiod when lap

conforms to this general pattern,general tothis conforms in 150g an in of experimental et al.et 2011;Prank et al. ) was the most most the ) was et al. 1998b). 1998b). et al. et al. et

Accepted Article for for pollinationAugust effective to from of October. end diameter rangesdiameter 18to 22 from grain Pollen size. & plant 2007), according Bretagnolle to (Fumanal, Chauvel grains plant per by pollen production varies wind. The among plants years 0.1pollen to3.8billion and from but release, directlyafter is sticky(Fig. 6C) only up to 6 hours. Although Although hours. to 6 only up su morningafter i.e. usually & Brush2010), in the 1965). American populations American flowers100% racemespistillate – eventheterminal consist only InNorthof females. have populations few European in a individuals unpublishedLeitsch-Vitalos, Some data). lateral branches of plants cutinSeptember develop almost exclusively female flowers (M. BasalMeade 1981). & (Barbour cutting (mid-August) late from branches male racemes than significantly more cutting earlyheads.Lateralfrom regrowth produces (mid-July) female develop to shoots tend bracts.lateral High-ranking small by surrounded groups(1–5(10)) heads are clustered in the axils of foliar leaves below the terminal male racemes in small 2011). Female Benoit & Simard Bretagnolle 2007; & Chauvel drymass(Fumanal, with plant according1015) canbevery but are theplantsize positively variable correlated to strongly (15–93,mean =55, per raceme = 142, mean (5–2878, Raceme number 1964). involucrum (Payne but elongatingin numerous dense and arranged Common ragweed is exclusively wind-pollinated. (A) FLORAL BIOLOGY VIII. Floral andseed characters This isarticle by protected copyright.reserved. Allrights observed across the native range (Ziska season pollen start ofthe inthe synchronisation of level remarkable responsible the for southernindividuals from populations & (Leiblein Tackenberg may 2014).This be seedsgrew anddispersed pollen flowered upto than and five and smaller weeks earlier experiment were highly with correlated latit garden common a growthand phenology late flowering in Forinstance, & Tackenberg 2014).

Anthers open with a rise in temperature and low relative humidity (Martin, Chamecki low (Martin, temperaturerelative humidity and withrise in opena Anthers c. 5% of the exclusively 5% individuals the have of flowers pistillate (Gebben Ambrosia artemisiifolia Ambrosia μ n m (Taramarcaz

= 1015), and florets per capitulum (9–39, capitulum per mean=18, florets = 1015),and et al. after some hours it is dry enough to be dispersed dry enough be is hoursit to after some 2011). ude, i.e. individuals northernfrom populations racemes. The phyllaries form a funnel-shaped et al. et nrise, nrise, and pollen release afrom flower lasts Male capitula are short-stalked (2–5 mm) (2–5 mm) areshort-stalked capitula Male exclusivelyis anemophilous,the pollen 2005).The pollen load ishigh enough

n = 203), capitulum = 203),capitulum number n = Accepted Article seeds. seeds. viable to produce hybridfails this However, 1958). Beals & (Wagner America North rarely in of Hybrids HYBRIDS (B) Shykoff(Genton, & Giraud 2005; Chun genetic data indicate heterozygosity deficits, probably through some degree of selfing 100%outcrossi for Crompton 1975)butalso This isarticle by protected copyright.reserved. Allrights in populations on arable is negativelyin populationson fields af Seed production seeds/plant;fields (c.2,300–6,000 2007). ChauvelFumanal, &Bretagnolle 300–2,500 c. productionaverage (between seed seeds;and Russia Fisjunov (62,000 ruderalhabitats Individuals lowerhad 1984). from of value extreme 94,900 seeds was also plant; G.counted Kazinczi, a data)for unpublished andinterspecific competitio intra- (without ofnumbers1971). However, inHungary plantbeen have seedsfound highest the reported per production,higher 3, seed ranging from rangehad in thenative Plantedindividuals 2007). Bretagnolle & Chauvel (Fumanal, plant (±271 2,518 of Franceshowed number anaverage SD)seedsperpopulations in ragweed seed are likely tobe the determinants the of seeds. number most of important A survey offive suitability 2007). habitat Therefore, Bretagnolle andcompetition1971; Fumanal, Chauvel & Seed productionof commonis ragweed closelyplant to biomass related (Dickerson & Sweet (C) SEEDPRODUCTION DISPERSAL AND A. artemisiifolia (Bassettsterile Crompton 1975).Nohybridshave& with hybrids created (1988) Lauzer &Cappadocia Gilles, 1963). (Steyermark America infrequently inNorth which been 2006), also Committee has found Cappadiocia (Vincent & Editorial 1988;FNA artemisiifolia & Swanton 2005; Nitzsche 2010). There is experimental evidence for some degree of self-pollination (Bassett (Bassett & degree self-pollination some of for evidence is experimental There Ambrosia ×helenae Ambrosia A. artemisiifolia and . A. psilostachya A. trifida Rouleau hybrid between a is with congeners have been reportedrarely. Hybrids between through cross-fertilization that resemble through cross-fertilization ( A. ×intergradiensA. et al. 135–32,485 seeds per plant (Dickerson & seeds Sweet perplant(Dickerson 135–32,485 n generally between 18,000 and but48,000, an 2010; Karrer 2010; Karrer fected by the densitybyfected crops the of (Chikoye, Weise ng (Friedman & Barrett 2008). Population &Barrett 2008).Population (Friedman ng seeds/plant) than individuals arable from seeds/plant) than yet been found intherange yet been non-native offound W. H. Wagner) have been observed observed been have Wagner) H. W.

et al. A. artemisiifolia A. 2011; Gaudeul A. trifida and et al. A. trifida A. 2011). but are A. A.

Accepted Article seed dispersal by human activities (Bullock dispersal byactivities seed human explain therates observed spreadat thelandsca cannotbyofand wind dissemination seeds gravity isobvious simple dispersal,distance it that also be induced by dark, low temperature fluctuations and high CO high fluctuationstemperature and low by dark, be induced also 1975; Bask 1972;Willemsen BazzazRice 1979; (Bazzaz dormancy the next Willemsen springgerminate entersecondary 1970; (induced) & around 4° fewdays a (optimally temperatures (inna primary This inautumn. dispersal their (Bazzazstrategies Bakker 1970;Thompson, Ambrosia artemisiifolia (D) VIABILITY OF SEEDS AND GERMINATION mother plant. with Dickerson (1968) who showed that disper 0.45 m and percentile ma 99% of1.05 (A. Lemke, plastic sheets around single of plants ra common (hydrochory) (Fumanal (Fumanal (hydrochory) dispersalseed by Zoochory2010). birds(Nitzsche of floating seeds dispersal by and water inEurope, although reports are there of spread on theone plants or for weeks two afterripening. (Béres 1981;G. pollination maturity following 4.65 mg 24populations for and China. inEurope Similarl Germany. in population experimental and Hunga Germany from populations several 3.60 mg (Fumanal (Fumanal summer periods. secondary1979). Under conditions, field dorma This isarticle by protected copyright.reserved. Allrights shown (Rosas While the achenes have no obvious morphological adaptations vectorsfor of long- Seed mass is highly variable within individual plants and differs between plants withinhighly individual Seed mass populations anddiffers is variable In North America, Gebben (1965) stated that Gebben(1965) stated America, In North A. artemisiifolia artemisiifolia A. et al.et et al. 2007). Mean seed mass in different populations in France ranged from 1.72 to et al. 2008). Bullock et al. et forms apersistent soilseedbank asaresult germination of complex 2007). Nitzsche (2010) found a seed mass (2010) mean a 2007).Nitzsche found around5mg for and dipersal (epizoochory) by animals ( animals by (epizoochory) dipersal and 2007) seems to have minor importance. et al . (2012)found littleevidence. for seed dispersal by et al.et & & Bekker 1997).Seeds ar of caching of ragweed seeds by rodents and of of seeds ragweed and byrodents caching of of y, y, of amean found (unpublisheddata) Karrer pe scale, which are probably achieved through through achieved probably are which scale, pe Karrer unpublished data). Seeds Seeds tend tostay unpublisheddata). Karrer te) dormancy is then broken by broken lowwinter then is te) dormancy C; Willemsen, C; 1975) and that seeds do not in & Baskin 1980). Secondary 1980). in& Baskin can dormancy sal bysal wind is restricted to2 m around the gweed gave amediandispersal distance of ry and ahigh mean 10 of mg for one Seeds take 4–6 about weeks to develop to ncy may also be induced during dry induced also be may hot ncy 2012). Dispersal2012). withsticky experiments unpublished data). This is in accordance seed dispersalby a plays birds role in the

Bison bison Bison 2 concentration (Bazzaz concentration e dormant following e dormant ) has also been ) has Accepted Article (1985) the dormancy period starts in May in the native region. & Baskin Baskin According to until mid-July. to 79% increases up ratethen germination and layer) soil cm the 0–20 seeds in dormant of May (2–6% broken until be could Hunyadi 1984).Fumanal, Gaudot &Bretagnolle conditions, seed dormancy January brokenfrom is toFebruary1975; (Willemsen Béres & strati some monthsbefore need toripen for after astheystratification their optimum germinate at donot seeds freshly harvesteddormant obtain 75% about (Baskin of germination & Baskin 1987; Fumanal (2013) showed reduceddelayed and germination w two-fold to twelve-fold higher when soil surface was disturbed. Furthermore, Fenesi seed bank in an invaded set-aside wasfield doubled when competitors wereremoved and a field experiment, Fumanal, Gaudot & Bretagnol (Baz competition lackof the and disturbance This isarticle by protected copyright.reserved. Allrights number of authors (Raynal & Bazzaz authors(Raynal 1973; number of within fiveyears seeds for stored in bags paper at room temperature(Kazinczi seeds quickly lose their overviability four buried seeds 22 cm germinated at did so respectively, germinated in vitro 30years after burial depth. Intotal, 57% 21% ofand seeds in buried the soil at and8 cm 22 cm, seedson their dependsconditions, thesurvival in of field &Baskin Likewise, (Baskin 1977). conditions experimental under specific in time germinate to inability ability or buttheir seeds germinate after 40years. Theseresults donecessarily not indicate life-spanthe maximum of able to still in1879were byinitiated Beal theexperiment seeds from ragweed of common Wa Brown & & 1946; Stoller decades (Toole for land) to 4477(inset-asideland) to habitats) seeds m inva different in seed-bank the (2008) studied CardinaWebster, Squiers & &Sheeley 1993; of early seedling and establishment Germination Because of secondary dormancy, stratification 4°Ctwo weeksto dark at by for bebroken wet, dormancy can Seed Seed-bank densities of Seed-bank A. artemisiifolia artemisiifolia A. A. artemisiifolia A. artemisiifolia fication (Brandes & Nitzsc & fication (Brandes after 39 years (Toole after 39years & Brown (Toole Unburied1946). years (Béres 2003).Viabil Bigwood & Inouye 1988; Gross 1990; Rothrock,InouyeBigwood Gross 1990; & 1988; zaz 1979;Rothrock, & InSquiers 1993). Sheeley - ² in the upper 20 cm of banksin upper20 soil. crop cm Seed in the ² ded habitats and found on average 536 (in waste 536 onaverage found and habitats ded White 2003). Fumanal, Gaudot & Bretagnolle Bretagnolle Gaudot 2003).Fumanal, & White ofin1902, and6% started Duvel’s experiment in in field-crophabitats been have analyzed by a le (2008) observed that recruitment from the recruitment that (2008) observed from le x 1974). Darlington (1922) showed that 4% (2008) observed inFrance dormancythat ith thepresenceith of adultplant competitors. A. artemisiifolia A.

seeds

can remain can alive in the soil he 2006). Undernatural 2006). he et al. et ity decreased by 73% are mostly related to related mostly are 2006). However, et al. et 2012). et al. et

Accepted Article potential from -0.8 to -1.28 MPa (Sartorato & Pignata 2008; Shresta Shresta 2008;Pignata & -1.28 (Sartorato -0.8 to MPa potential from °C water temperature minimum to ranges germination 3.6 3.4 The minimum from for and (Pickett & Baskin 1973; Guillemin & Chauvel 2011), or high salinity (DiTommaso 2004). This isarticle by protected copyright.reserved. Allrights CO changes, humidity suchas soil and temperature light, of various temperature, rhythm factors, Chauvel & 2008) ornot(Guillemin Bretagnolle germinationis type, habitat 1975; 2011). & Chauvel Stoller & Guillemin onthe 1968; Depending Wax 1965; Dickerson seeds on the soil surface and decreases with increasing depth, ceasing below 8 cm (Gebben of soildisturbance. of distribution 585, = mean layer, soil mean deeper layer, = 1066; (upper soil disturbed less patternwasobservedopposite for =1630, (mean 592, =(mean (0–5cm) upper soillayer waslower inthe seeds andset-aside fieldsof extant werelarger Thenumber in habitats. ruderal than conditions (Martinez (Martinez conditions artemisiifolia Therefore, germination). MPa(no until-1.6 potential osmotic increased linearly with al. Guillemin 1993; 2003; CardinaWebster, &White Fumanal, Gaudot2008). &Bretagnolle habitat disturbanceamount of and (Forcella to the seeds thefrom uppersoil arerelativelylayers – low andvariable 2.3from to 42% – according Bretagnolle (Fumanal,spring Gaudot 2008),recruitment & early ofgerminate buried rates in day/8 In hnightphotoperiod. conditions, natural if even most ofthe seeds are toable °C a16h °C 25/20 with 15/10 seeds stratified at and90%at of germination observed 80% 1980; Shrestha Baskin Baskin & °C (Shrestha (Shrestha °C temperature,germination ofstratified seeds in 2013). However, according to Sang 2 (Bazzaz1970;concentration 1973;Raynal &Baskin 1968, Pickett &Bazzaz 1973; Under experimental conditions (after stratification), seed germination is highest for for highest isgermination seed (after stratification), conditions experimental Under Germination is delayed Germination is bydelayed & (Pickett Baskin temperatures low 1973; Willemsen 1975; et al. can germinate under a very high wate et al. A. artemisiifolia 2013), low (Shrestha moisture n = 240) for frequently ploughed habitats such as crop fields, whereasthe crop fields, as such habitats frequentlyploughed for = 240) 1999; Sang et al. 2002). positively correlated positively with seed correlated Gaudot (Fumanal, mass & seed bank isspatially aggregated seed bank irrespectiveof theintensity et al.et Ambrosia artemisiifolia Ambrosia et al. 2001; Nitzsche 2010). Guillemin & Chauvel (2011) 2010). Guillemin&Chauvel 2001; Nitzsche et al. 1999). Undercontrolled conditions and constantat (2011) the germination of seeds decreases near- decreases seeds of germination the (2011) creases from 5 to 25 °C and decreases up to 40decreases upto 25°C 5to from creases and et al. conditions (wastelandand set-aside habitats) 2011). Germination in the field depends field inthe on2011). Germination n et al. = 240) than inthe (5–15cm)240) than = layer deeper 1999;Guillemin r deficit as well as under water-logged as r deficit aswell under

1992; Rothrock, Sheeley & Squiers is also able to germinate is alsoable to distilled in n et al. = 250). The horizontal The = 250). et al. et 1999; Guillemin 1999;Guillemin 2013),low light A. A. et Accepted Article which haswhich been found on only2011). The exception the is nearly were all recorded in theformer Yugoslavia, Hungary and Russia In Eurasia,some 50insect andone acarine species havebeen on found OR PARASITES (A) FEEDERS IX. Herbivory and disease and epicotyl areglabrous an is correlate positively size Seedling germination. minutely hairy (Kazinczi photosynthetically active (Bazzaz 1973). The primar surface. They become green shortly they after emerge the from soil,to be andstart often the that margins to lower the extends purple of show pigmentation a hairless and isGermination epigeal (Dickerson 1968; Fig. 7). The cotyledons elliptic, are short-stalked, (E) SEEDLINGMORPHOLOGY than field crop populations and werevery competitive in such environments. road populations saline, growing in observed that germinate (from 5–12%) at high NaClconcentrationsmmol L 5–12%) at (from 400 (i.e. germinate This isarticle by protected copyright.reserved. Allrights L mmol 200 about of al. between (Sang pH5tooptimum pH at a maximum rateof germination 8and 98.2% 5.57 anwith 48%germination) least (at to 12 pH-values, 4 from of range large over a water Italy (Lombardia, Piemonte and Emilia-Romagna; Fig.and S2;Müller-Schärer Piemonte Italy (Lombardia, Eur time in thefirst detected for been 8) has wilt and virus, spotted tomato of the vectors Thysanopteraspeciesseveral including Rédei In Hungary Koczor (andprobably& 2008). elsewhere), with dominated survey 2011). InHungary, the on fauna in a hemipterans to little damage polyphagousthese species are andcause this species, whichwas accidentally introduced Despite its recent arrival in Europe, data from the rapidly infestedexpanding area suggest that artemisiifolia A. 2011). high ge isalso It able tomaintain Eupteryx atropunctata Eupteryx (Jenser, Kiss & 2009).Recently, Takács the beetle -1 of of NaCl (Sang et al. et A. artemisiifolia artemisiifolia A. d often purplishd often in colour. and Thefirstfoliar 2008a). leaf appears within few days after Emelyanoviana mollicula mollicula Emelyanoviana et al. Ostrinia orientalis Ostrinia Frankliniella occidentalis 2011; Eom, &WestonDiTommaso 2011;Eom, 2013) can and rmination rates (70–80%) under moderate salinity undermoderate (70–80%) rates rmination ope in southern Switzerland (Ticino) and northern northern and Switzerland (Ticino) southern ope in in China (Wan in China (Wan T. tabaci T. d with the weight theseed. Thehypocotylof into China and now used there as a most as there now used and China into -side habitats show hi y leaves areovate in outline, pinnate and has been shownto transmit this virus to

Mutuura Munroe (Crambidae), Mutuura & being most often recorded (Kiss, (Kiss, recorded often being most et al. A. artemisiifolia A. A. artemisiifolia A. 2003). 2003). ofHowever, many and Ophraella communa -1

gher germination gher rates germination (Table 2; Gerber Gerber 2; (Table Thrips tabaci ). DiTommaso (2004) DiTommaso ). A. artemisiifolia A. A. artemisiifolia artemisiifolia A. (Gerber (Gerber et al.et is a host to that are 2014). et al.et (Fig. et al. and et et

Accepted Article herbivoresthe inof specialized introduced range (Colautti (Genton folivores invasion. initsSimilarly, range invaded thisplant been shownhas tohave escaped insect populations of species, this linking me possible releas enemy that they proposed pathogens, predation, of levels soil folivory, pathogensreduced perhaps seed losses toseedand MacKay & &Kotanen KoMacDonald (2008) and that common ragweed escapecan natural enemies by dispersing to new sites. Although foli invertebrate by damage of levels reduced littleexperienced by isolated 100m existing common from as as ragweed of populations so only visit flowers to eat herbivores occur only 2).herbivores lowdensities at (Table occur pests,most native and includingknown agricultural somespecies, polyphagous mainly of (2008). In thecontrast, insect complex associated with ( snails Moreover, seed predation by ground beetles ( Moreover, predationground seed beetles by be specialists, associated plants with some areto likely 109species which of Palmer1995), (Goeden& America North andSouth herbivores ha asmanyas 450 total, up to 70species of insects and mites associated with This isarticle by protected copyright.reserved. Allrights theof genus most of these herbivores are leaf-chewing insects, the plant is also attacked by stem-galling artemisiifolia (Zhou agent biocontrol successful management by example for thebeetle herbivores, Trichia striolata

MacKay & Kotanen (2008) showed that in the native range, nativeexperimental populations range, the in (2008)showedthat Kotanen MacKay & In its native range, native range, In its ) and several polyphagous consumers substantially. et al. Epiblema 2005), consistent with the ReleaseEnemyHypothesis, the i.e. absence :

Gastropoda, Hygromiidae) has been reported by MacKay & Kotanen A. artemisiifolia (MacKay & Kotanen 2008). Goeden & Palmer (1995) identified Palmer (1995) & Goeden 2008). & Kotanen (MacKay the pollen (Bassett & Crompton 1975). et al. ve ve been found onthevarious native Ambrosiinae thesubtribe from (Gerber 2010), reduces growth and seed production of of production growth2010), reducesand seed is is attacked variety invertebrateby awide of vores seedand predators. These results indicate Zygogramma suturalis e can occur for native as well asexotic aswell native occurfor e can Harpalus chanisms of localspread with long-distance Ambrosia artemisiifolia Ambrosia including beetles, bugs and moths tanen (2010) found only slight effects of (2010)foundonly slight effects tanen A. artemisiifolia

A. artemisiifolia spp.; Coleoptera, Carabidae) spp.; and et al. 2004). (see (see in the native range. In is wind-pollinated, is in Europe consists Europe in Ambrosia spp. XI. et al. et

Biological . 2011). While in A. A. Accepted Article Crompton 1975).Ofthe 20 fungal the native range, numerous fungal pathogens are associated with campestris This isarticle by protected copyright.reserved. Allrights in the field (Kiss Gerber 2; (Table unpublished data), & Crompton1975; Krumbiegel 2007) as well as in Europe Follak and G. (S. Karrer artemisiifolia Ambrosia AND DISEASES C) PLANT PARASITES (B, events, e.g. the import of grain, contaminated oi we and in cities beenmade have populations, al. DiCeccoMandrioli, 1998), Russia & Andina (1865; http://www.ambrosia.ch/vorkommen-pflan (1883; Pyšek, Sádlo & Mandák Chauvel Ascherson 1874;Brandes2007;& Nitzsche collected thefor second half of the 19th century inmost countries: Germany (1860; summarizing its historywas by invasion early publication first the Europe, Eastern and ForCentral 2014). Crocker & (Lockton in 1836 wild, the Strasbourg). In Montpellier, 19th century gardens inat botanical least (Alençon, five another Angers, Avignon, 1763;Paris,1775;Poiti (Lyons, the 18thcentury France in that (2012) report artemisiifolia Ambrosia INTRODUCTION AND HISTORICAL SPREAD History X. Vajna 2002). Peronosporales), were recordedin Hungary and2002 in1999 (Vajna, Bohár & Kiss 2000; and Phyllachorales) fungal pathogens, 2010). Most early records of of records early Most et al. et ), which is also native to North America and has been introduced into Europe. In In Europe. into introduced been has and America North to native also is which ), 2006), Austria (1883; Essl, Dullinger Kleinbauer& 2009),Czech Republic et al. et al.et A. artemisiifolia Phyllachora ambrosiae 2011) most have 2011) a most widehosthave range littleand on impact theplant is only rarely attacked by plant parasites. In North America (Bassett (Bassett North America In parasites. by plant attacked is onlyrarely Plasmopara halstedii recordedwas inEurope inbotanicalgardens.first Bullock 2003). Outbreaks of diseaseof Outbreaks byepidemics caused biotrophic 2003). two A. artemisiifolia A. artemisiifolia 2002), Hungary (1888;Kazinczi sometimesis attacked by a parasitic dodder ( pathogens associatedpathogens with A. artemisiifolia wasleast present three inat gardensbotanical in Poppendieck 2007), France (1863; Bonnot1967; 2007), France Poppendieck (Berk. & M.A. Curtis) Sacc. (Acomycota: (Acomycota: Sacc. Curtis) &M.A. (Berk. (1918;CABI2014), Ro inEurope to pertain which ephemeral casual (Farl.) Berl. & De Toni (Oomycota: & De Toni(Oomycota: Berl. (Farl.) l-seed and seeds of forageof speciesNorthseeds from l-seed and re probably related to repeated introduction repeated to related re probably Ascherson (1874). First records have been been have First records (1874). Ascherson ze/vorkommen-von-ambr ers, 1791) and during the first half the half of first ers, the 1791)andduring was first was recorded asincasual a Britain

Ambrosia Ambrosia et al. mania (1907;Csontos mania 2008a), Switzerland 2008a),Switzerland species in Eurasia Eurasia in species species (Bassett & osia/), Italy (1902; osia/), Italy (1902; Cuscuta Cuscuta et al.et et et

Accepted Article large populationslarge in emerge fields (Novak regions, climatically most favourable inthe populations; naturalized increase, especiallyof numberlimited ofnaturalizedpopulations gain importance. artemisiifolia increase inconsiderably, particular in th Csontos regions (e.g.Kazinczi climatically 1920s the mostfavourable from in the 2009), Essl,fe railways; Dullinger &Kleinbauer Aust(e.g. in and repeatedintroductions dispersal smallof scattered populations;spreadmedi is Essl, Dullinger & Kleinbauer 2009; Csontos (Rybnicek (Rybnicek invasion can currently be classified as the phase ofincreased spread andnaturalization the delayed, inthe e.g. Germany, British Isles, Czech andPoland, Republic roads. along major often regions, climaticallyfavourable into expanding populations less large with records, numbers of in Central distinct invasionEasternand Europe four stagesreveals (Kazinczi apparently not been reached yet. with Hulme(Pyšek the 2005)–has, & heavilyexception the of most regions, invaded phasethe saturation of theinvasion – i.e.when rate the ofinvasion newareas slows down invasion Apost foci emerged. in areas hoc Europe first analysis of populations naturalized and first associatedthe populations localexpansion of to adjacent This isarticle by protected copyright.reserved. Allrights America III. Increased spread and naturalization 1960–c. and(c. spread 1990) naturalization Increased III. II. Incipient spread naturalizationand local (c. 1930–c.1960) century–c. (19th I) 1930) Rareintroductions IV. Rapid spread and increasing abundance (c. 1990–ongoing) abundance(c. andincreasing spread Rapid IV. In climatically lesssuitable regions, wheretheoccurrence invasion of these phases is ( et al. Brandes & Brandes & Nitzsche 2006; Chauvel et al. continues to be rare; reproduction and local spread from an increasing, yet still yet increasing, from spread an reproductionandlocal torare; be continues 2010). 2000; Brandes & Nitzsche 2006; Tokarska-Gudzik Nitzsche 2006; & Tokarska-Gudzik 2000;Brandes A. artemisiifolia A. isnaturalized Europe, increasingly in regions in larger et al. et e Pannonian Basin; but in most countries, most in but Basin; Pannonian e et al. et al. 2009); local spreadis becoming dominant. ated largely by anthropogenic long-distance largelyated by anthropogenic ria, 80% of early records are associated are earlywith records of ria, 80% 2010) (Fig. S3): w first naturalized populations are recorded naturalized populations are w first : thefew records: of 2006). Only2006). of occurrence the the after

A. artemisiifolia A. et al. et : a great increase: agreat in the : numbers of of records : numbers : numbers of of records : numbers A. artemisiifolia A. 2011). InEurope, 2011). A. artemisiifolia A. et al. et al. invasion invasion 2008a; 2008a; are A. A.

Accepted Article car at 30 mph (48 km h mph (48 car at30 to fosterlong-distanceas highdispersal as al Distances bridged by this arevector higher th Lippe von der 2009; & Karrer (Vitalos of frequencies small rather suggest vehicles 2011). Chauvel 1998; Prots & is relatively sparse(Song contamination of on frequency and levels but information fodder), andembankments, animal slopes cont other dispersedthrough is also species the that likely is However, it decreasing to be seems pathwaythis currently of importance containing wholegrain and seeds has recent seedsHowever, legislationof of anEU-wide restricting theamount This pathway ismainly res bird wasthat oftenfeed with significantcontaminated quantitiesof seeds of hasfood been investigatedin several studiesBullock (EFSA 2010; common ragweed seeds attached to roadside mowing machinery in infestedareas. 53.1aof seeds of mean report (2009) & Karrer by Vitalosdispersal mowingmachinery. Lippe der (von This isarticle by protected copyright.reserved. Allrights and France (Chauvel railways) (Essl,Dullinger asobserved in e.g. network (roadsides, 2009) Austria Kleinbauer & importance and spatial range (Table 3; Bullock artemisiifolia (see (barochory) plant parent the of metres) mechanisms (Bullock aswell whereas local population growth andshort-distan long-distanceThe medium- spread of and PATHWAYS The spread of spread The

Common ragweed is toable disperse quickly and efficiently along the transport can be dispersed be by relativecan differing human waysactivities in many with et al. 2013). Another vector that could explain rapid spread along roadsides is rapid spreadalong roadsides explain could that 2013).Another vector A. artemisiifolia A. et al. -1 ) showed a median distance) showeda of one 99% ofmeter anda 9m quantile 2006). However, experimental quantification of dispersal by dispersal experimental quantification of 2006).However, et al. ponsible for introductions into private gardens andparks. gardens introductionsinto private ponsible for 2012). Seeds of through commercial bird feed and small domesticpet feed through commercial bird and et al.et ong roads. Dispersal kernels after 80 passes of of 80passes onga roads. Dispersal kernels after an primary bydispersal gravity and wind but not ly been adopted (EU Subsequently,2012). the A. artemisiifolia aminated commodities (e.g.aminated commodities for seedmixtures Seed Production and Dispersal seedsby transported windslip of the vehicles et al. et al. 2013; Milakovic, Fiedler& Karrer Milakovic, 2013; ce spreaddependent are dispersal on natural A. artemisiifolia A. 2012).

is driven by human agency, et al. et et al. stay close (within a few stayclose a (within A. artemisiifolia A. 2012). They showed 2012). 2004, Karrer 2004, A. artemisiifolia. ). However, However, ).

in food food in 2014a). et al. et A. A.

Accepted Article ranges. SDMs constructed todepictwere the distribution Guisan species (SDMs; models & Thuiller 2005) to be andfitted projected across negligible expansion and limited unfilling; Petitpierre Petitpierre unfilling; limited expansion negligible and has remained mostlyin climat analogue stable Guisan Petitpierre by reported 2014).(Petitpierre This isarticle by protected copyright.reserved. Allrights range shifts, range shifts, tolatitudinal In all England.in but southeast addition limiting factors climateremain may With regards to Britain, summers cool and commonragweed’s preference for a continental (e.g. southern infested RussiaandUkraine). arecurrently drought that of because inregions al. expect Drought particular, high in temperatures. become climatically unsuitable to due the comb will range inEurope distribution currently parts southernmost 2014). Incontrast, the of of spread in the harvestersof severaldozens up to31 133seeds perin harvester eastern Austria. Karrer example, artemisiifolia Scandinavia and theBritish (CunzeIsles southern asfar northas climatically 21stcenturyin Europe regions bythe mid suitable change climatewarmer scenarios,medium delayed expandthe summersfrosts and later summers For spread. willautumninstance, frosts and allow undernorthward anduphill changeto climate (Table There 4). consensus is among that warmer continental-scale models spread due orpotential future common ragweed’s predict modelling studies European Several SPREAD FUTURE landfill sites (Rich 1994) as one of the habitats invaded by Consequently, several authors listconstruction 2013), may limit for common ragweed. Indeed, some models predict population declines some predict models population Indeed, ragweed. for common may 2013), limit Agricultural machinery strongly is implicated the in oftransport seeds of Using the distribution of distribution Using the The transportation of soil, gravel, constructi transportation of The et al. et 2014) wereconducted, which revealed theclimaticthat niche of A. artemisiifoliaA. within and within between fields (Taramarcaz et al. A. artemisiifolia A. (2011) heavy found varying of but of levels contamination soybean et al. et is predicted also to higher invade mountainelevations in areas (2012), detailed (2012), niche conservatismclimatic analyses (see A. artemisiifolia A. as well (Taramarcazas well et al. et es between (i.e. nativeand non-nativeranges its es ined effect of increasing summer droughts and current andfuturecurrent distributions based on the ed to strengthen in southern Europe (Jacob southernto strengthenEurope (Jacob in ed 2013; Chapman Chapman 2013; (e.g. Essl, Dullinger &Kleinbauer 2009)or in North America, Eurasia and Australia andAustralia Eurasia inNorth America, on material and landfill waste is involved waste involved is andlandfill material on et al. et al.

et al. A. artemisiifolia 2005;Chauvel 2012; Appendix S1), allowing 2012;Appendix 2005; Bullock et al. 2014; Storkey . et al. A. artemisiifolia A. et al. 2006).For 2012). et al. A. A. et et

Accepted Article niche, i.e. that populations of This isarticle by protected copyright.reserved. Allrights reconstructing the most likely introduction pathways 2011; (Gentilini Bullock and allows predictions suitability improve environmental hasshownto modelling with dispersal coupling spreading, still is species the where in countries Finally, models. yethave Europe to inthe inincluded be useherbicidesuch as greater northwest and fertiliser of have operating models well-mapped beencountries in spread fit able dynamics time seriesto pathwaysdispersal models 2012). inadequate for Onlyregional-scaleanthropogenic (Bullock, sizes in and variation population theat spreadof continental poor scale, mapping of mapping is by hampered Modelling 2010). S5). (Fig. Eurasia rangein theinvaded of front north-eastern onthe occur latter cells mainly These S5). predicted to become suitable in a warmer climate in 2100(potential range expansion, Fig. are predicted become to (potential unsuitable rangewhereas constriction), 5023grid cells are only1579 km) currently suitable, = 8151change. (resolution c.50 0.5°, Amongthe gridcells the potentialdistribution increasewill – globally country suitable that most the is SDMs Oceania. New Finally, and Japan. Zealand in predict of S rim southern inSouthAmerica, the Chile Brazil, Paraguay, species Argentina Southern the (Fig. Uruguay, and for 9): highly suitable of distribution (all other between known 0.07 S4).Outside 0.2; variables Fig. ofthe (0.814) than and aboutevaluationAnnual indices).information of distribution potential depictthe ability to species’current climatic 9;see details). (Fig. Appendixniche S1 for Theof SDMs’ evaluation invasion (Bullock as deliberate eradication or enhanced phytosanitary regulation of seed the trade may restrict such efforts Control decline. followedabandonment succession bylonger-term be likely will artemisiifolia abandonment Eastern(Spangenberg in Europe A. artemisiifolia Finally, all the models assume temporal and spatialassume conservatismof the and ecological the temporal models all Finally, However,there uncertainty is predicting in spread of the any invasivespecies(Gallien, Several factors beyond climate may also influence future spread. Ongoing agricultural spread. Ongoingagricultural influence beyondclimate future also may factors Several ’s invasion’s (Bullock A. artemisiifolia etal. ’s distribution (Richter’s distribution 2012; Richter Richter 2012; A. artemisiifoliaA. used to calibrate SDMs, several areas are predicted to be etal. A. artemisiifolia A. et al 2012). Proliferation in the earlyin the stages post- 2012). Proliferation of et al. . 2013a,b). Other important effectsanthropogenic Other. 2013a,b). important A. artemisiifolia artemisiifolia A. 2013a,b). have retained the same the have retained outh Africa, and, in Asia, in outh Africa, and, China,SouthKorea et al. and also onthe British– under Isles climate mean temperature is far more important far is temperature mean ’s non-equilibrium distribution, poor ’s non-equilibrium distribution, 2012) may profoundly 2012)may influence

is excellent(see Apendix S1 for climatic tolerances in tolerances climatic et al. 2012). A. A. Accepted Article Corre & BretagnolleCorre & additive genetic may variance and havepromoted rapid adaptation (Chun,and evolution Le This populations admixture introduced Europe. mayof increased genetic have diversity and vectorsseparate introductions of two source inNorthAmerica from areas occurred have in 2005; Giraud Gaudeul & Shykoff by distance ofisolation significant pattern was Easternand West twogeographical in the at least Martin America. in North origin (Gaudeul the 20 middleof populations established the in of the century at and introduced the populationsend 19th Western Europe between from Geneticof genetic exist differences diversity. high range, leadingits non-native levels to of studies introductions suggested multiple common ragweed of countries within parts in most was observed in China (Liwas inChina observed bottleneck events associated withtheinitial introduction of the species whereas no such effect expansionofGiraud (2005),range 2005; Fumanal2007; Chun populationsand non-native overall showsimilar genetic Thislittle differentiation. holdsas in EuropeAmerica, well and North as bothnative between indicates which occurs within populations, ratherthan observed variation genetic This isarticle by protected copyright.reserved. Allrights populations 2005; &(Genton, Shykoff Giraud Chun VARIATION GENETIC invasion. of modelling predictive into areas currently or toocold dry for in (e.g.scale habitat level), its range may ragweed ongoing adaptation allow to extend common detected the at of level the realized macro-climatic niche (Petitpierre (Genton (Leiblein- germination oftolerance its frost placesall times. Evidence ofat hasbeen duringall evolutionary invasion in found adaptation

Introduced populations in Europe and Asia in populations and Europe Introduced et al. et et al. 2005; 2005; Chun 2011;Gladieux

2011). et al. et al. et et al. et al. 2012). Nuclear chloroplast mi and et al.et 2010; Martin 2010;Martin 2011; Hodgins &Rieseberg but2011), this has not been et al.et 2011), which could be regionsof by explained different 2011),whichcould A. artemisiifolia (2014) found weak but significant isolation bybut significant distance isolation found weak (2014) 2011). A hypothesis for the European is thatEuropean findings the A 2011). for hypothesis Wild, Wild, &TackenbergKaviani 2014) phenology and vasion, presenting a significant challenge for detected along an Eastdetectedalong an to cline(Genton,West et al. et genetic diversity (Genton, Shykoff & Giraud &Giraud Shykoff genetic(Genton, diversity ern clusters in Europe. In North America, InNorthAmerica, a Europe. in ern clusters th century in Central and Eastern Europe andEastern Central in century th are probably a mixture of different native different of are probablymixture a 2014).According toGenton, Shykoff & occurred in France after aseries of

et al. 2010;Li crosatellitesused inthese etal. et al. 2012). Atfiner 2012). 2012). Most Accepted Article sugar beet, oil-pumpkin, potatoes, various legumes andvegetables. soybean crops like sunflower,sown maize followedand row areby mostheavily invaded Delabays 2006) and recently from Geeastern (Rhône Valley) (Chauvel be heavily infested. In Western Europe, large populations in fields are known France from ha on which700,000 Hungary, recorded millionhectares out of to 5.4 considered in waswere weed abundance in winter wheat and maize weed fields inwinterwheatand abundance in Hungary (Novak populations populations andmay modify theragweedof responsecommon toabiotic conditions. genes a proportionof maydifferentially be countries. native areas(Chunfrom active gene flow between inco olderpopulations, suggestsand genetic This populationsthat ones. diversityhave older than current arisen from specimens from the 19th and 20th centuries. Recent populations in France show greater allelic become a dominant weed dominant become inarable fields a in andEasternEurope, In Central Agriculture IMPACTS XI. Conservation This isarticle by protected copyright.reserved. Allrights 2009; Galzina 2009; Galzina Chun results of The allocation. reproductive greater evolvedhave a on pressures Riesberg suggest 2011).Theseresults abio that observedreproduction in thesame population sample in previousa study (Hodgins & stress responses.be therefore may They involved increasein growth an of rate and genes, several are potentially involved inthemetabolism of secondarycompounds and in Amongthose conditions. expressed differentlygenes 180candidate these identified under populationsintroduced subjected to stress.light ornutrient This genome-wide approach 45,000 native between than genes and of more expression Riesbergby comparing the (2011) from 21st (1950), to8th 21st(1950), from Changes ofgenetic diversity have reconstructedover time usingbeen herbarium The molecular basis The molecular of invasiveness of A. artemisiifolia A. et al. et , Impacts and Management and Management Impacts 2010; Follak & Fertsak 2012). For &Fertsak instance, 2010; Follak

(1970), to 4th (1970), to et al. et al. populations, such that populations at or higher latitudesaltitudes 2010) and also from crop seedsEuropean traded crop 2010) andalso between from 2006), locally from Switzerland (Bohren, Mermillod 2006), locallySwitzerland from & particular in the Pannonian Plain, Plain, Pannonian particular in the

(1988) to 1st (1996-97; 2007-08) (1988) to1st(1996-97; (Týr, Vereš &Lacko-Bartošová(Týr, Novak 2009; rmany (Schröder& Meinlschmid 2009). Spring- A. artemisiifolia A. rporating new alleles from newintroductions rporatingfrom new alleles expressed within native within expressed non-native and tic conditions might have exerted selection exerted might conditions have tic

was studied by Hodgins & was byHodgins& studied A. artemisiifolia A. et al. et al. (2011) indicated that (2011) indicated A. artemisiifolia in place in terms of place in 2009). In 2003 it increased et al.et has has

Accepted Article types (Dahl, Strandhede & Wihl 1999 and references therein). Northern andHemisphere) induces as reportedly inthe latetoOctober August andautumn (typically symptomsfrom summer in causes Ambrosia artemisiifolia Health 20% and80% in 1991 and1993,respectively. maximumloss the varied density, soybean yield between it in was 70%andinmaize 65 and competitive with maize and soybean crops (Canada).in Ontario For high (Smith ranges introduced 2003)and Bernstein & (White native its in allergy of pollen major cause become a plantsof four per10mrow. Weaver of &Ritter(1981)Williams calculatedyield8% soybean of at an losses differing clim because of situation European to butthesewith caution 2003), results can only betransferred Cowbrough, Brown&Tardif low canopyheight andwhen Swanton 1995; Cowbrough, Brown2003). Damage &Tardif is especially high in crops with a densities. In sugar beet, plots infested with infested plots beet, Insugar densities. This isarticle by protected copyright.reserved. Allrights More data are available from North American studies 2000). Inthesestudies, considerable cropyield lo resulted inan average lossof of sugar 50% yield compared to weed-free plots (Bosak &Mod 2002; Varga 2002; Varga has been(Bosakmaize2000; Varga, by documented &Mod &Reisinger Beres authors some impact of various densities of relative tothecrop, and density of the et al. in so be (particularly substantial can loss Yield its to contribute cutting regenerate after to plants m plants Kazinczi example, 2009),butdependsthe type, largely oncrop of the time emergence of

The species’ temporal emergence pattern, pattern, emergence species’ temporal The -2 caused in 33%yield reduction 21and sunflower 30%almost maizeand atboth in etal. et al. 2013). 2006; Kazinczi 2006; et al. et is a noxious plant that has highly allergenic pollen (Fig. 10), (Fig. pollen which has 10), that highlyis anoxiousplant allergenic (2007) demonstrated that (2007)demonstrated A. artemisiifolia A. A. artemisiifolia A. et al. et al. 2007; Nitzsche 2010; Bullock 2010; 2007;Nitzsche A. artemisiifolia A. success as a weed (Bassett & Crompton 1975). A. artemisiifolia A. emerges together with the crop. In Europe, the (2001) also found (2001) found also atic conditions and cropping practices. Coble, andcroppingconditions atic practices. low-growing such as crops beets;Buttenschøn on the yield of sunflower, sugar beet and and beet sugar sunflower, of yield the on rapid and plastic growth, and strong ability plastic ability andstrong growth, and rapid thma twice thma oftenabout as as other pollen sses atlow occurredeven weed densities. A. artemisiifolia

(e.g. Coble, 1981; Williams& Ritter

infestation (Chikoye, Weise Weise infestation(Chikoye, & at a density of2 Ambrosia artemisiifolia Ambrosia A. artemisiifolia atdensities of 5and10 A. artemisiifolia et al. A. artemisiifolia A. A. artemisiifolia to 2012). For For 2012). 5 plants m 5 plants was highly highly was density density has -2

Accepted Article to induce symptoms may be very low,e.g. 1–3 pollen grains m to of sensitisation relevance Clinical over the short(Tosi over term the isexposedto increasesthe it human population of time with incidence allergyin a Consequently, (Asero2007). publichealth has recentlya seriousfor problem asthe become Theprevalence2009a). of to sensitization Europe was 10.7% (ranging from in Finland1.4% to49.7% in Hungary; (Burbach the that showed (3034patients), countries consideredand is tobe specific for agood marker Gleich 1978; Gadermaier & Goodfriend in allergen major the represents itsuch as and proteins, is which 38-kDa non-glycosylateda protein that belongs tothe family pectatelyaseof against Ambrosia clinical relevance because sufferers of with as well where the plant is not frequently found or absent (Stach orabsent not found frequently is theplant where of transport long distance pollen it is not possible to ithigh theconsequences isnot ascertain of pollen (Burbach been shown to increase Amb a 1 expression (Singer This isarticle by protected copyright.reserved. Allrights 1988), but rangethe typical 5–20pollen is grains m environmental conditions such as increased concentrations of atmospheric CO of atmospheric concentrations increased as conditions such environmental (Wopfner v 1from Art as well a5from Amb 1and species distribution (Amba allergen restricted groupswith et al. Šikoparija 2013). The allergenic capacityThe allergenic of 2013). 2010). Cross-reactive It has been suggested that the atmospheric concentrations of The majority (>90%) of (>90%)of majority The Allergic cross-reactivity is Allergic cross-reactivity freque Artemisia Artemisia etal. pollen pollen allergens (Taramaracaz et al. et al. et Artemisia 2009b), 2009b), e.g. asdocumentedItaly where Northern for 2009, 2013) 2009, a also is concern for allergy (Bullock 2005). Theallergenicity of was also detected species (White & Bernstein 2003). The closelygenera related have Artemisia etal. Ambrosia Ambrosia 2011). Ambrosia ) and pan-allergens (profilins, polcalcins, and the nsLTPs) the and polcalcins, (profilins, pan-allergens ) and Ambrosia

in 31% of ragweed-sensitized 31% ofin patientsin Hungary (Páldy pollen from from centres of pollen Ambrosia Ambrosia and et al. et ntbetween species within the genus Artemisia et al Ambrosia clinically relevant sensitisation rate rate to sensitisation clinically relevant pollen-allergic patients are to sensitized Ambapollen-allergic patients 1, pollen pollen transported over long distances remains Artemisia . 2008; Gadermaier, Hauser 2013) &Ferreira 2005). For 2005). E(IgE) Immunoglobulin instance, Ambrosia pollen allergen determined in 14 European 14 in determined allergen pollen Ambrosia et al. pollen allergypollen reactfrequently to also pollen has increased over time in Europe Europe in overtime hasincreased pollen -3 (Bullock (Bullock pollen allergens divided canbe into

2005). pollen can change underdifferent et al. et sensitization sensitization (Smith Ambrosia Ambrosia Ambrosia et al. 2007;Smith -3 d Ambrosia et al. et Ambrosia -1 2012). As a result, the distribution into areas distribution pollen concentrations concentrations pollen (Comtois (Comtois & Gagnon pollen (Adolphson,pollen 2012; Prank pollen required pollen allergy allergy pollen 2 et al. etal. Ambrosia which have have which Ambrosia Ambrosia Ambrosia Ambrosia 2013). 2008; et al. et et al. as in as

Accepted Article can be best achieved bysieving techniques. containments mu and exported imported Accordingly, This isarticle by protected copyright.reserved. Allrights artemisiifolia amount of allowed sincelimits the 2012.This regulationmaximum EU in the been inplace limiting for contam the standards Strict pathways. desi be to have Preventivemeasures populations. prevention of dispersalanthropogenic seed an of management The physical Physical management MANAGEMENT The synthesis habitatof of affiliation Nature conservation sensitized individuals. al. containing uncertain (Cecchi (since 1 January 2013; EU 2012) to 50mg kg January 1(since EU2012)to 2013; plant species and thus have an indirect nature conservation impact (Bullock Blackburn artemisiifolia ephemeral. the Accordingly, and are densities (e.g. value herb opentall communities, forests). Usually, these populations occurin low al. nodiscernible negativefar onthe has been (Fried invaded impact identified communities PannonianBasin(Bullock most invaded, inthe regularly rivers are grazing)created have of that Similarly, soil. open open patches and sand gravel banks along seem butthese mostlyShevera Mosyakin 2006), & Protopopova, Botta-Dukát2004; Ukraine: 2006;Hungary: &Klingenstein & Nawrath Mihály Countries(Austria:Essl, F. Ka G. European Occurrences dryin grasslands have been documented several times andEastern for Central occurrences have been inthe recorded centre ofits current distribution Europe. in highsuch value only of andmostof it that habitats invades nature reveals conservation rarely 2014).Common ragweed occassionallymay habitats other colonize of high conservation 2013) indicates that thesepollengrains that in 2013)indicates thepotential reactions to allergic have induce Ambrosia et al.et seeds in bird seeds (since 1 January2012; in Europe qualify as ‘no impact’ according to the impact according‘no impact’ assessmentqualify the inof Europe as to scheme (2014). Management measures (2014).Management against et al. et pollencollected duringepisodes of transportlong distance (Grewling 2010), but a recent study showing th A. artemisiifolia A. A. artemisiifolia A. Ambrosia rrer, unpublished data; Germany: Alberternst, Germany: Alberternst, data; unpublished rrer, environmental consequences of invasion of of invasion of consequences environmental d mechanical control of already established ofestablished already control mechanical d can have twocomplementary have approaches: can gned specifically for the the introduction various for specifically gned -1 ination of feed stuff by seedshave offeedstuff ragweed ination , which is , which 10–12seeds.equivalent to c. to be a consequence ofbe a disturbancesto (e.g. st now be kept almost free of seeds; this

EU 2011) and in animal material feed animal and in EU 2011) artemisiifolia in Europe by Bullock inEurope by Bullock

A. artemisiifolia A. e presence of Amb a 1 in samples1 in ofpresence Amb a e et al. seed loads on roadside roadside on loads seed 2012). so However, may affect otheraffect may et al. et al. 2012). (2012) et A. A. A. A. et et Accepted Article that develop to additional to develop enable branches that as close to the soil surface as possible (Bohr This isarticle by protected copyright.reserved. Allrights (Karrer resprouts from seeds every Subsequent cuts 3weeks & Karrer 2014b). shouldfirst cut bedone be weeks the after 2–3 the effectively, set however, seed reduce To production. to minimize pollen optimal 2008c) is conditions of southern Central Europe,M whereas cutting before onset the of flowering toend (mid of July under the climatic pollen,racemes andof released male number reductionsinthe to onlytall) moderate leads cm plants20 arec. cutting that (2011) showed (when early VidottoFerrero Patracchini, & 2011). Meiss 1981;Bohren,& Meade Delabays 2006;Bohren,Mermillod Mermillod &Delabays 2008; potential from below buds cutting very high (Bassett & is height Crompton 1975; Barbour & contact dermatitis. the onset gloves flowering andtowear of(male) Karrer treatmentsdestroyor heatingburningcomposters; germinationability (e.g. that in requires the seeds ripened with plants Pulling of Delabays 2006). & Mermillod hand-pulling (Bohren, also but effective, most the Therefore, 2010). (Nitzsche not prevent reproduction todelay of flowering initiation butdoes reported the 2010; Nitzsche 2010;Patracchini, Vidotto &Ferrero The cutting 2011). vegetativeof is plants Benoit & Simard & Aarssen1996; stems lateral (Irwin growth existing of increase base or the le stem apex and the suchof as removaldamage et al. (Kazincziavailable arecontrol pathway. this concerning regulations legal with country European new infections (U.Starfinger cleaned (Karrer avoid thetransport of seeds from infested fiel thou several tensof reach can data) unpublished (Vitalos 2009) cutting machines &Karrer 2011; Bullock et al. et al. et Mowing is themostwidely applied mechanical technique.control As the regrowth Once common ragweed inaregion, several established is mechanical techniques for 2008; Karrer 2008;Karrer 2011) and it must be done carefully to avoid seed losses.be donecarefully before avoid is act seed 2011)anditmust to It bestto et al. 2011). Transport of 2011).Transportof contaminated soil et al et al. . 2012). However, et al. et al. et 2011), to 2011), cutting multiplication branch avoid bedone should et al.et 2011;Karrer & Pixner 2012). 2008c; Buttenschøn, Waldispühl & Bohren 2009; Karrer unpublished Currently, data). Switzerland is onlythe ilakovic, ilakovic, &KarrerFiedler 2014a; Kazinczi en, Mermillod & Mermillod en, Delabays 2006).Accessory buds prolongation of seasonal growth (Karrer seasonal growth (Karrer of prolongation ds and roadsides, machinery must be machinerydsthoroughly must be and roadsides, A. artemisiifolia ginning of male flowering (Milakovic, Fiedler ginningflowering (Milakovic, male of and crop harvesting crop (G.Karrerand machinery very optionis mechanical control laborious, aves, as the plants regenerate from buds aves, from plantsregenerate as the from sand seeds per machine. Thus, in order to order in Thus, machine. seeds per sand in order to prevent skin irritations due due skinto inorderto prevent irritations are important of important production to preventare the

is an additional serious source for source serious additional is an tolerates substantial physical substantial tolerates et al. et et al. et

Accepted Article some non-target species. be techniquesSuch appliedtosmall can ragweed (Buttenschøn, Waldispühl & Bohren 2009; U. Starfinger (Karrer technique this of applicability the limit dates germination in plasticity and seed bank soil persistant al. experiment, be effective inof densities reducing has been suggested as a method control (Béres 2004; Kazinczi years (Swanton bank within seed few soil of the depletion to the leads spring in period germination the duringweeks 4–6 grubbing every ploughing or Shallow & Delabays cropharvest(Bohren, Mermillod be by2008). ploughing achieved after early Vladimirov & Maneva 2010) &Maneva Vladimirov seed showedidentical (Karrer mixtures restoration with results was rapidly (Meiss outcompeted planting that combined experiments This isarticle by protected copyright.reserved. Allrights perenne of 2009).Control &Bohren (Buttenschøn, Waldispühl mechanical hoeing techniques have been shown to reduce ragweed densities in crops Schröder & Meinlschmid 2009; Meinlschmid & Schröder crops) a limited number less of he effective vegetables,various legume oil-pumpkin, (e.g. crops minor herbicides, in emergence while crops, major In USA. the 2,4-D in to control inherbicidesagriculture years, widely used For 50 been have Chemical management artemisiifolia 2010). 2006; Karrer Establishing a closed vegetation cover in thermaltreatments steam,Experimental (hot that includingrotation, susceptibleare less crop crops Introducing to Tillage (ploughing, grubbing, hoeing) may kill kill may hoeing) grubbing, (ploughing, Tillage (by more than 95%). (by 95%). more than Lolium perenne , reducing its biomass by 91%. et al. et al. 2011). 2011). and and .

Ambrosia artemisiifolia

Buttenschøn,Chauvel& Bohren 2009;Gauvrit & Waldispühl Dactylis glomerata Medicago sativa et al. Medicago sativa Medicago A. artemisiifolia A. A. artemisiifolia A. artemisiifolia 2008; Meiss 2010).2008; Meiss A. artemisiifolia rbicides can be applied (e.g. Kazinczi(e.g. applied canbe rbicides combination with mowinghasshowncombination with been to also also showed an inhibitory effect on outcompeted outcompeted with intensive cutting, cutting, intensive with can be can controlled with pre- andpost- flaming)been control have found to biomass was reduced most by biomass wasreduced most A. artemisiifolia artemisiifolia A.

(MacDonald & Kotanen 2010). In & (MacDonald 2010). Kotanen populations, and they may also kill A. artemisiifolia artemisiifolia A. Ambrosia artemisiifolia Ambrosia et al. et al. et A. artemisiifolia A. et al. 2011). Inaglasshouse 2011). 2008c). However, 2008c). the et al. on can fields stubble A. artemisiifolia unpublished unpublished data). 2000; Murphy A. artemisiifolia,A. A. artemisiifolia A. plants. Various et al. (Milanova, (Milanova, 2008c; grown grown Lolium Lolium e.g. A. A. et

Accepted Article herbicides that target the enzyme acetolactate resistance seemsisolated, tobe risk the that resistant ecotype was Hungaryfound in 2009). Although& Taller Cernak (Cseh, of case this 2009). crops in Hungary. crops in fields & (Schröder 2009; Kukorelli Meinlschmid herbicidescertain (e.g. imazamox, tribenuron-methyl)an mightinfested bealternative for application isgreatly sunflower reduced. Thecultivation that of cultivars are tolerant to is which theAsteraceaeparticularly herbicide insunflower, potential in also of the andthus been observedAmerica North (Patzoldt in decades many cases of resistance of observed between triazine and substituted urea herbicides (Heap 2014). During thelast two Saint-Louis, (linuron; infesting carrots in theinfesting maize USAStephenson (atrazine; ragweed in observed was photosynthesis inhibiting toresistance some herbicides first, ragweed have been detected in various crops herbicide-resistant ecotypes.development of This isarticle by protected copyright.reserved. Allrights application methods, which are recommended for the control of of control for the arewhich recommended application methods, an exhaustiveof(2008c) andKazinczi list active provide &Novák and ingredients (2012) Kazinczi less infested. (e.g. are cereals) densities with high plant crops as autumn-sown Meinlschmid 2009). Moreover, herbicide applica & Schröder Bohren 2009; & (Buttenschøn, Waldispühl period has along germination species because recommendedlastingis the achieve activeingredientsleaf control, a and soil to of may efficiency combination herbicide Sequential treatments anda often survive. improve Ambrosia artemisiifolia influenced by theplant growth stage at application Mermillod (Bohren, & Delabays 2008). impacts on other plant species. The efficacy of several active ingredients against speciesthe is of production and seed active ingredients such as glyphosate and glufosinate are to appropriate control bothpollen In Europe, as of a its intensive sinceIn Europe, the 1960s, atrazine- consequence application an Yield losses are most important in certain The dependence upon herbicides of for control A. artemisiifolia A. is most susceptible at the 2–4 leaf stage, while larger individuals individuals whilelarger stage, the2–4 leaf at susceptible is most A. artemisiifoliaA. (Gauvrit & Chauvel 2010), but (Gauvrit Chauvel 2010), they strong have & also DiTommaso &Watson 2005).Cross-resistanceDiTommaso was common ragweed populations ragweed common resistant become to et al. sincethemid-1970s Canada USA. the and At in Herbicide-resistant populations of common Herbicide-resistant synthase (ALS) is because of important the 2001; Taylor Taylor 2001; et al. tion should be with combined crop rotation springcrops (sorghum, soybean) andmore et al. toALS andglyphosate have inhibitors 2011). In non-crop areas, non-selective 2011).In non-crop areas, A. artemisiifolia

1990) and in Canada in ragweed in Canada in 1990)and et al. et A. artemisiifolia 2002; Brewer&Oliver in fields has led to the in main in et al.et

Accepted Article This isarticle by protected copyright.reserved. Allrights (Stojanovi candefacta T. recently, More an (Kovalev agent inEurope invasive 1971a). against control plant non-native biological a introduction of intentional first 1969wasthe region inRussia Krasnodar for classicalthe of biological control on damage reach natural enemies native very Europe, few Europe (Gerber Teshler 1988; & Templeton range (Cartwright Common ragweed has been a target for biological management both in itsof parts native managementBiological diversifying methods. control Gauvritby & and (Grangeot, 2010) Chauvel ingredients alternating byactive on roadsides avoid managers isto land inEurope and famers in the cropping system to ragweed common reduce density. Therefore, a objective of major if rapidly to resistance may lead rotation cycle intensive generally, the whole orinMore mixtures) useofinthe crop ALS (alone inhibitors sunflower varieties (Chauvel &Gard2010). tolerant inhibitor increasing ofALS cultivation artemisiifolia densities 5,000 high as densities m as beetle Russia in with this obtained the results Yugoslavia in 1985andagain (nowCroatia) in 1990(Igrc,DeLoach Zlof & Atfirst, 1995). 1998). Griffiths & (Julien Kazakhstan released Kazakhstan,Ukrain in and Georgia wasGriffithsthealsospecies North 1998). sameyear, In & in the Caucasus the (Julien and 2005has promotedthe build-up oflarge populations. thefrom harsh continental climate in the region, but a series of mild winters between 2002 and therelease of the North American noctuid moth artemisiifolia layeggs,emerge and adults of sincedamage inspring when period short occur during the 1999). However, population outbreaks anddestruct In 1978, the leaf beetle leaf the In 1978, ć

A. artemisiifolia et al. plants, thereby increasing crop yield cropyield plants, thereby increasing plantsoverareas (e.g. duetoherbivory large of has also been recorded in the Ukraine (Poltavsky & Artokhin 2006) and Serbiabeenrecorded has the 2006) Artokhin also in Ukraine (Poltavsky & et al 2011). According toPoltavsky this &Artokhin(2006), moth has suffered . 2011), Australia (Palmer (Palmer 2011), Australia . . Therefore,using specialist naturalenemies from range native the -2 Zygogramma suturalis in an arable field in southern Russia anarable southern fieldall anddestroyed in in A. artemisiifolia A. Zygogramma suturalis were very promising (Reznik 1991). It reached very(Reznik 1991). were promising integrated cultural practices are not introduced et al. e, but establishmentis e, in but onlyconfirmed et al the selection of resistant plants in fields infields and plants of resistant selection the high enough densities to inflict significant toinflict densities high enough 2002) as well as inthe invadedrange in . 2010) and Asia (Zhou. 2010)andAsia ion of hostplant populationsion canonly F. was released and quickly established quickly released was and F. two- to threefold (Goeden &Andres (Goeden two- tothreefold in Europe was initiated in the 1960s, inthe Europewasinitiated in

Zygogramma suturalis Zygogramma suturalis

candefacta was released in formerwas released Huebner inthe Huebner et al . 2010). In larvae) A. A. A. A.

Accepted Article most most highwith populations enough defoliate completely flowering to and prevent andseedset of This isarticle by protected copyright.reserved. Allrights diapauseresult can insummer in provokes and female oviposition inhibition FA1209 ‘Alien Challenge’ and FA1203 ‘Sustainable management of management Challenge’FA1209 andFA1203 ‘Sustainable ‘Alien ecologydistribution, and morphology. FE ac knowledgeon colleagues whoshared their to numerous We are obliged Acknowledgements areas in Europe with warm summer temperatures (Zhou Europe significantly, but due toits preferences climatic Thus, this biological may agent control be ab re in growingseason results during the generations used used asuccessfulas biological agent against control (Müller-Schärer Switzerland Italy andsouthern innorthern found damage have declined strongly since (Julien & Griffiths 1998). of infection is achieved (Hartmann & Watson 1980). In Russia, be very damaging andsignificantly reducespollen andseed production if systemic infection 1998). Attackby Griffiths & (Julien USSR the former Canadafrom into Albugo tragopogonis pathogen the early 1960s, Inthe Europe. biological organisms been control elsewhereagents into have usedas accidentallyintroduced (Reznik was negligible weed target average densities Russia consequently, population in very impactwere and, the low on the surveysthat 2006Databetweenfield from indicated (Reznik 1991). 2005and conducted ATOPICA project (Grant # 282687). The work of several authors has benefitted from the has workof authors several The benefitted (Grant #282687). ATOPICAEC project EUFP7 the with collaboration also benefited This from article (SMARTER). in Europe’ By theendof 2013, Airport, suggesting that International 2011). Firstobservations of A. artemisiifolia A. In 2013, the leaf beetle the leaf In 2013, control biological agents, twoexotic releasesof these deliberate to In addition A. artemisiifolia A. O. communa plants (Müller-Schärer (Müller-Schärer plants (D.C.) Gray) (Oomycota: Albuginaceae) was accidentally introduced O. communa and reduction in biomass and seed production seed biomass and in reduction and Ophraella communa etal. had already an 20,000area of colonized c. km O. communaO. 2007). in Italy in made werein thearea of Malpensa Milano Pustula tragopogonis Pustula et al. knowledges support from EU COST Actions Actions EU COST support from knowledges le toreduce ragweed common infestations in 2014). Population build-up due to multiple to due build-up Population 2014). peated and extended attack of single plants. may have been introduced by traffic. introduced been may air have Le Sage (Chrysomelidae; Fig. 8) was was 8) Fig. (Chrysomelidae; Sage Le P. tragopogonis A. artemisiifolia A. O. communa et al.et

2010). et al. (Pers.) Thines (synonym Thines (Pers.) may remain restricted to mayrestricted remain Ambrosia artemisiifolia initially caused heavy heavy caused initially in China in(Guo China 2014). This beetle is 2014).This beetle P. tragopogonis P. A. artemisiifolia A. , but levels of of levels but Z. suturalis Z. 2 (Fig. S2),(Fig. et al. can can

Accepted Article Barbour, B. & Meade, J.A. (1981) Meade, J.A. B. & Barbour, Asero, R. (2007) The changing pattern of ra (2007)The of changingR. pattern Asero, Ascherson, P. (1874) Ascherson, Anton, Zuloaga, A.M. & Republica vascularesde la Plantas Argentina. (2014) Flora F.O. in parts ofthe (1943) Theoccurrence ragweedsand H. other A. North American their Allard, Amano, K. (1986)Host range and geographical relations of water to the Data L.(1976) Almádi, Klingens S.& Nawrath, Alberternst, B., allergens withragweed (1978) Reactivity of G.J. Gleich, L. & Goodfriend, C.,Adolphson, This isarticle by protected copyright.reserved. Allrights Acevedo-Rodríguez, P. Strong, & M.T. (2007) References Davy. A.J. Editor, the and greatly benefited from comments of M.C,F. D.T. Proctor, Streeter, P.A. Thomas, M. Usher has publication This reproducing11. 10and for Figures Research) (Weed Wiley obliged to Excellence Program'. For thepermission to reproduce drawings of frameworkthe SocialFund in European G. Kazinczi bythe wassupported HALT (07.0322/2010/586340/SUB/B2). AMBROSIA’ Ambrosia – onmethods inEurope the invasion research ‘Complex to halt project DG ENV Weed Science Society Science Weed common ragweed 62 Europa. Europa. Naturschutzsicht. Argentina. URLhttp://www.floraargentina.edu.ar. Accessed 13 January 2014. Japan. Tokyo, Press, Society Science Japan elatior world. Einschleppungswege von 197–210. cross-reactivity. of determination and Analyses the antibodies: IgE byleukocytehistamine radioallergosorbentand release test 2014. URL Indies. http://botany.si.edu/antilles/WestIn , 1097–1099. vízáztartásához). Science, Botanische Zeitung Botanische

98, Nachrichtenblatt des Deutschen Pflanzenschutzdiensts Ambrosia artemisiifolia Ambrosia 292–294. Ambrosia artemisiifolia Ambrosia , 85 Botanikai KözleményekBotanikai , 82–86. Ambrosia artemisiifolia , 48 , 770–773. The effect of cutting height effect of of The and on anthesis date Journal of Allergy and Clinical Immunologyand Clinical Journal of Allergy gweed allergy in the area of Milan, Italy. gweed Italy. areaofallergy Milan, the in of TÁMOP-4.2.4.A/2-11/1-2012-0001 'National of TÁMOP-4.2.4.A/2-11/1-2012-0001 (Asteraceae). tein, F. (2006) Biologie, Verbreitung und Verbreitung (2006) Biologie, F. tein, , bisherein , nicht beachteter Einwanderer in Catalogue of the seed theseed theCatalogue of West of plants distribution of powdery of the fungi. distribution mildew , 66 dies/catalog.htm. Accesseddies/catalog.htm. 13 January Ambrosia elatior , 199–204. inDeutschland und ausBewertung

Proceedings of the Northeasternof the Proceedings A. artemisiifolia (Adatok az , 58 , 279–285. Ambrosia Ambrosia Allergy we are , 62 , , Accepted Article Béres, I. (1994) New investigations on (1994) NewBéres, I.the biology investigations of This isarticle by protected copyright.reserved. Allrights ragweed ( common biology of importance and (2003) Distribution, I. Béres, ecology succession. physiologicalThe plant (1979) Bazzaz, F.A. of Bazzaz,of F.A. (1974)Ecophysiology of Bazzaz, F.A.(1973) Photosynthesis Bazzaz,Secondary (1970) of the in seedsragweed dormancy the F.A. common Bazzaz, F.A.Succession (1968) onabandoned Southern Hills, inthe Shawnee Illinois.fields Bassett, Crompton,I.J. & (1975) C.W. Baskin, &Baskin, J.M. (1980)Ecophysiology C.C. seeds secondary dormancyof in of Baskin,C.C. Baskin, & (1985) The annua J.M. Baskin, J.M. & Baskin, requirements for C.C.after- (1987)Temperature ripening inburied Béres, I. weed(2004) Integrated management strategies of Belcher, E. (1985) Basset, I.J & Crompton, C.W. (1982)Th Baskin, & J.M. te Baskin, (1977) Roleof C.C. Basky, Z. (2009) Effect of native aphid species on the development of invasive ragweed of development aphidon the species native of (2009) Effect Basky, Z. Universiteit Gent, Universiteit FaculteitMededelingen enToegepasteLandbouwkundige Biologische Wetenschappen, artemisiifolia stratégiák). ürömlevel andSystematicsEcology Ecology artemisiifolia Ecology L.. artemisiifolia Ambrosia continuum. weeds.ofsummer annual seeds four and in the field. summer annual weeds. Atlanta. Ambrosia artemisiifolia Canadian Journalof Science Plant , , 55 49

ű artemisiifolia parlagf Magyar GyomkutatásMagyar és Technológia , 112–119. , 924–936. BioScience L.). L.). . L. and Bulletin of the Torrey Botanical Club Botanical Torrey of the Bulletin Handbook on Seeds of Browse-Shrubs and Forbs Seeds Browse-Shrubs and of on Handbook American Midland Naturalist Növényvédelem 59 ű A. psilostachya A. ( , 1295–1297. Béres, I. (1981)biology Distribution, of , Oecologia . (L.) in Hungary. Ambrosia artemisiifolia 35 , Ecology 10 , 492–498. , 351–371. , , , 61 30 39 DC. DC. Ambrosia , 475–480. Weed Research , 377–382. , 293–302. , Ambrosia artemisiifolia Ambrosia e biology Canadian weeds. of 55. The biology of Canadian biology Canadian The 11. of Weeds. 62 Redia Canadian Journal of Plant Science , 1003–1010. mperature in the germination ecology thegermination threemperature of in ,

artemisiifolia 90 l dormancy cycle l dormancy cycle in buriedweed seeds: a , XCII integrált elleni gyomszabályozásiL.) , , 186–190. 1 , 3–14. , , 97 27 , 211–213.

, 302–305. , 385–389. L. plantsgrown greenhouse in Ambrosia artemisiifolia Ambrosia artemisiifolia Ambrosia : a successional: a dominant. . USA Forest Service, Service, Forest USA . Annual Review of Review Annual Ambrosia trifida , 55 , 463–476. Ambrosia Ambrosia Ambrosia Ambrosia Ambrosia (Az L. L.

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Accepted Article Brückner, D.J.,Brückner, of Lepossa, A. ( (2003)& Herpai,Z. ragweed Inhibitory effect CABI (Centre for Agricultural Bioscience Agricultural 2014)for International) CABI Invasive Species (Centre This isarticle by protected copyright.reserved. Allrights Brouillet, L., Coursol, F., Meades, S.J., Favreau, M., Anions, M., Bélisle, P. & Desmet, P. Burbach, G.J., Heinzerling,L.M., Edenharter, G., Bachert, C.,Bindslev-Jensen, C., Bonini, Bullock, J., Chapman, D., Schaffer, S., Roy, Girardello,D., M., Haynes, T., Beal, S., Brewer, &Oliver, ( C.E. L.R. Braun, U. (1995) &Nitzsche,Brandes, D. (2007)Verbreitung, und Soziologie Ökologie von J. Buttenschøn, R.M., Waldispühl, S. & (2009)GuidelinesBohren, C. for management of G.J.,Burbach, Heinzerling, L.M., Rohnelt,C., K.C.,Behrendt, Bergmann, &Zuberbier, H. T. hypochondriacus artemisiifolia URL http://data.canadensys.net/vascan.Accessed 13January 2014. VASCAN,(2014) la données basede vasculairesdes plantes duCanada. 573. common ( ragweed resistant artemisiifolia Deutschen Pflanzenschutzdiensts ULR Accessed http://www.euphresco.org. 13January 2014. effects of common ragweed in Europe (E Europe in ragweed common of effects (2012) C., S. &Brough, Finch, M., Skinner, Hilbert, A., Schrauwen, A., Prank, M., Sofiev, M., Niemelä, S., Räisänen, P., Lees, B., Wheeler, B., Dickie, I., Phang, Z., Tinch, R., Civic, K., Delbaere, B., Jones-Walters, L., Jena, Germany. relevance ofrelevance inhalant allergen sensitizations in Europe. (2009a) study clinical GA2LEN II: T. skintest Zuberbier, A., Wöhrl, S. & Wiesner, M.,Orosz, Papadopoulos, N. G., Röhnelt, C., Stingl, G., A., Todo-Bom, E., VonMutius, Gjomarkaj, M., Gramiccioni, C., Haahtela, T., Kowalski, M.L., Magyar, P.,Muraközi, G., Canonica, G.W., Darsow, U.,Demoly, P., Durham, S., Fokkens, W.J., Giavi, S., Bousquet, J.,S., Bousquet-Rouanet, Bousquet, L., P.J., Bresciani, Bruno, M., A., Report. Final prevalence. increasingsuggests study GA(2)LEN - Europe in sensitization Ragweed (2009b) common ragweed, Allergy L. in Mitteleuropa. L.)-inflorescence extract on the germination of germinationof on the extract L.)-inflorescence ThePowdery mildews (Erysiphales)Europe. of L. and oftwosoil growth algae. , Ambrosia artemisiifolia Ambrosia 64, 664–665. 2009) Confirmation resistance mechanisms and glyphosate- in Ambrosia artemisiifoliaAmbrosia , Tuexenia 58 , 286–291. NV.B2/ETU/2010/0037). European Commission, European NV.B2/ETU/2010/0037). , . EUPHRESCO project AMBROSIA. EUPHRESCO 2008 project 27 Assessing and controlling the spread and Assessingcontrolling spread the and the , 167–194. Chemosphere ) in Arkansas.

Allergy , 64 , 51 , 1507–1515. Weed Science Weed Gustav Fischer Verlag, Verlag, Fischer Gustav , 515–519. Amaranthus , Ambrosia Ambrosia 57 , 567– , ‐ 09.

Accepted Article Colautti, R.I., Ricciardi, A., Grigorovich, I. Grigorovich, Ricciardi,A., R.I., Colautti, H.D., &Coble, F.M. Common (1981). ragweedWilliams, ( Ritter, R.L. This isarticle by protected copyright.reserved. Allrights Chauvel, Dessaint,B., F., Cardinal-Legrand, C. & Bretagnolle, F. (2006) historical The Chun, Y.J., Bret Fumanal,B.B., & Laitung, Chun, Y.J., Bretagnolle,F.(2011) Le V. & Corre, ragweed of D.,Chikoye, ( common C.J. &Swanton, (1995)Influence S.F. Weise, Chauvel, B. & B.Gard, à (2010)Gérerl’ambroisie d’armoise. feuilles of limitations (1988)Biological G.E. Templeton, & Cartwright, R.D. Comtois, P. & Gagnon,(1988) Concentratio L. & P. Comtois, Cecchi, L., Testi, S., Campi, P. & Orlandini, Chapman, D.S., Haynes, D., Beal, S., Essl, F. Chauvel, B., Vieren, E., Fumanal, B. &Bretag explained byexplained the Enemy ReleaseHypothesis ? artemisiifolia artemisiifolia astheadmixture primary post-inva Biogeography of spread 144. Accessed 13January 2014. http://www.cabi.org/isc/?compid=5&dsid Compendium: des Végétaux des a mycoherbicide forgiant ragweed, pollinose: uneméthode pour déterm pollinose: 20 trait amonginvasive Science artemisiifoli pollen doespollen notinducenew sensitizations in the shortterm. d'Allergologie et d'Immunologie Clinique native and invasive range limits of common ragweed. ragweed. common of limits range invasive and native Colloque international sur la biologie des mauvaises herbes artemisiifolia d'Ambrosia 202. , 1378–1388. , 43 , 375–380. Ambrosia artemisiifolia a) time a) time of emergence and density on beanwhite ( , ) interference in soybean ( soybean in interference ) ) populations in France. , 633 33 , 665–673. , 12–16. Ambrosia artemisiifolia Ambrosia artemisiifolia Ambrosia L. via les semences de tournesol. tournesol. de semences les via L.

Ambrosia trifida L.records.in France herbarium from New Phytologist sion processesin ragweedcommon ( Glycine max , =4691&loadmodule=datasheet&page=481&site= A. & MacIsaac, H.J. (2004)Is invasion success iner les seuils cliniques. cliniques. inerles seuils 28 agnolle, agnolle, F. (2010) Gene flow andpopulation S. (2010) Long-distance transport of ragweed of transport Long-distance S. (2010) , 279–286. & Bullock, J. (2014) Phenology predicts the Phenology (2014) predicts J. & Bullock, n pollinique et fréquence des symptômes de etfréquence des npollinique nolle, F. dedissemination(2004) Possibilite F. nolle, populations in France. Ecology Adaptive divergence forafitness-related divergence Adaptive ). . Plant Disease , Weed Science Weed

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15, Weed Science 293–297. Ambrosia artemisiifolia Ambrosia Ambrosia artemisiifolia in Central and Eastern Europe. ,

46 ogical characters,ogical encroaching strategy andcontrol habit of (2009) Molecular characterization of (2009) Molecularatrazine resistance characterization in , 555–560. L.) phenological development and growth. growth. and development phenological L.) Acta Agriculturae Shanghai Weed Science Influences ofInfluences mycorrhizae and phosphorus on L.). L.). Ambrosia artemisiifolia L.), PhD thesis, Cornell University, Ann University, Ann Cornell L.), PhD thesis, Influence of temperature, photoperiod, and photoperiod, of temperature, Influence Journal ofAppliedGeneticsJournal (exceptdistribution France) before 2009. ,

52 Botanica Helvetica ISRN Ecology , 1002–1009. Environmental Experimental , Weed Science Weed 16 Weed Science Weed Ambrosia artemisiifolia Ambrosia L. an invasive weedin , 73–77. American Journal ofAmerican Journal , 2013 , , 120 , ID610126. 50 , , 321–327. , 75–78. 19 , Ambrosia Ambrosia Ambrosia Ambrosia Ambrosia 51 , 64–66. , 947– Weed Weed ) Accepted Article EFSA (European Food Safety Authority) FoodSafety (2010)Sc EFSA (European Fisher, N.H. & Quijano, L. (1985) Allelopathic agents from common weeds. from agents (1985) Allelopathic L. Quijano, Fisher,& N.H. recognition kin Plant (2012) S.A. Dudley, & H. Maherali, J., Klironomos, A.L., File, Re Commission Union)(2012) EU (European Re Commission (2011) Union) EU (European FNA (Flora North America) (2006) Volume 21: Magnoliophyta:VolumeFNA 21: part): (2006) NorthAmerica) (Flora (in Asteridae This isarticle by protected copyright.reserved. Allrights Fenesi, A., & E. Albert,Á.-J. Ruprecht, ability (2014) predictfuture Fine-tuned to Essl, F., Dullinger, S. & Kleinbauer, I. (2009) Changes inthespatio-temporal patterns and Pr Plant Mediterranean and EPPO (European Eom, S.H., DiTommaso, A. &Weston, L.A. of (2013) soilsalinityEffects growth the of in (1992) D.Paulissen, W. & Werner, R., V., Wirth, Düll, Weber, H.E., H., Ellenberg, &Hayova,Dudka, I.A. (2007) V.P. animal health or on the oron health environment onthethe presenceof animal of seeds feed. in Ukraine. palmeri enhances abundanceof partner. symbiotic microbial in environment competitive dioxins. dioxins, asregardsCouncil maximum levels for arse theof Parliament and European the of 2002/32/EC Directive II to I and amending Annexes oftheJournal European Union cons and histomonostats and coccidiostats certain of carry-over spp. and Ambrosia melamine, nitrite, levels for maximum regards as amending Annex I to Directive 2002/32/EC of th 119–133. of preferences habitat http://www.eppo.int. ULR 5.3.1. Version Nutrition Plant artemisiifolia Ambrosia vonMitteleuropa. in Pflanzen Zeigerwerte Symposium Series,Washington.Symposium Plants between InteractionsBiochemical ESFA Journal , Official Journal of the European Union Ambrosia Ambrosia artemisiifolia Mikologiya i Fitopatologiya , 36 , 2191–2204. spp., diclazuril and lasalocid andA sodium action thresholds for , 8 , 1566. Ambrosia artemisiifolia Ambrosia biotypes collectedfrom roadside andagricultural field. Ambrosia artemisiifolia , L159 Plasmopara angustiterminalis Plasmopara , and related weeds. , 7–24. ,

41 Accessed 15 April 2013. Scripta Geobotanica gulation (EU) No 744/2012 of 16 August 2012 16August 2012 of gulation (EU)No744/2012 , 12–19. (ed Thompson),A.C. 133–147. ACS pp. olidating Annexes I and II thereto. thereto. II Iand olidating Annexes gulation (EU) No 574/2011of 16 June2011 otection Organization) (2013) PQR database during its invasion in in Austria. during invasion its , nic, fluorine, lead, mercury, endosulfan, mercury,nic, lead, fluorine, e European Parliament and of the Councilthe and Parliament European e of ientific opinion onientific or theeffectpublic on L219 seeds. seeds. PLoS One

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17 (Asteraceae). in der Landwirtschaft in Österreich. Österreich. in derLandwirtschaft in , 25–35. , 48 , 349–359. Ambrosia artemisiifolia Biological Invasions, Biological L. L. Ambrosia artemisiifolia Ambrosia , 100 Seed Science Research Science Seed Annals ofAnnals Botany Weed Science Weed Preslia , 305–313. L. PhD L. Thesis,University ofBurgundy, Dijon. Annals ofAgricultural andEnvironmental Medicine , Ambrosia trifida,Ambrosia Artemisia annua, Iva xanthiifolia 85 zur und Verbreitung Bekämpfung zur der , 41–61. , 2007) Estimation of pollen pollen 2007)and of production Estimation seed 40 etagnolle, F. (2006) Which role can (2006) Which F. can arbuscularetagnolle, role le, Chauvel,F. & large Can the B. (2008) seeds: What consequences for itsinvasion in Ambrosia artemisiifolia , DOI 10.1007/s10530-013-0597-6 DOI 10.1007/s10530-013-0597-6 ive processesive ofan invasive species plant in environmental control of environmental andtemporal control size- , 636–644. D. & F.Essl, (2013)Invasion dynamics of 101 Seed-bank dynamics in the invasive plant, explain its invasive success in France? invasivesuccess explainits in , , 1303–1309. 18 Evolution 0–1. , 101–114.

Proceedings of the 67thALVA- the of Proceedings , 65 , 2061–2074. L. invasion in France? in invasion L. Ambrosia Ambrosia Ambrosia , 14 ) , Accepted Article Gauvrit, &Chauvel, Gauvrit, C. B. (2010)Sensitivity of Gadermaier, G., Wopfner, N., Wallner, M., Egger, Didierlaurent, M., A., Regl, G., Aberger, Gladieux, P., Giraud, T., Kiss, L. & Genton, B. (2011) Distinct invasion sources of common invasion sources Distinct P., T.,Gladieux, Giraud, B. (2011) Genton, Kiss, L.& This isarticle by protected copyright.reserved. Allrights Genton, B.J.,(2005) T. J. Shykoff, A.&Giraud, ( ragweedThe Gebben, common ecology A.I. (1965) of and chloroplast Nuclear (2011) J.A. Shykoff, & L. Kiss, T, Giraud, M, Gaudeul, Gard, B., Bretagnolle, F., Dessaint F. & Laitung Boul C.H., Albert, Münkemüller, T., L., Gallien, Genton, B.J., Kotanen, P.M., Cheptou, P.O., Adolphe, C. & Shykoff, J.A. (2005) Enemy (2005) J.A. & Shykoff, Cheptou,C.Adolphe, P.O., P.M., B.J., Kotanen, Genton, Bari N., Galzina, Gilles, V.,D. Lauzer, Cappadocia, M. (1988)& Schaffner, E., Gerber, Gassmann, U., A., M. Hinz, Müller-Schärer, Seier, (2011)H. H.L., & glyphosate at various developmental stages. pollen allergens. mugwort Lang, F. &Hawranek, R., F., Ferreira, T. recombinant andnatural molecules. ragweed ( 26 ragweed, the invasionintroductions multiple in history show commonmicrosatellites worldwide of 28–35. ragweed common exhibit strongtolerance to foliar damage. 75 weed invasive Distributions where here? togo invasive species: of from distributions potential of introduction. ofragweed, common populations easternMichigan. PhDthesis, University of Michigan,Ann Arbor. reciprocal transplantreciprocal experiment. nobut release evolutionary defence loss of in aplant invasion: an inter-continental Common Ragweed,Common past. biological for Prospects of control , 574–576. , 75–81. Weed Research Weed Ambrosia artemisiifolia Ambrosia artemisiifolia Ambrosia , 16 Ambrosia artemisiifolia ć Molecular Ecology , K., Š , 331–342. Ambrosia , 51 ć , 559–573. epanovi Allergy

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Biological Invasions Biological to glufosinate and and glufosinate to . Diversity and Weed Science L.) in south- L.)in , 14 , , , , Accepted Article Goeden,R.D. Lessons Palmer, (1995) & W.A. lear Guisan, A. spec W. (2005) Predicting & Thuiller Guisan, A.,B.,B., Petitpierre, Broennimann, Daehler,C. (2014)Unifying &Kueffer, C. S.,Reib J.P., A., Granger, Guillemin, Gardarin, of Effects (2011) & Chauvel, B. J.-P. Guillemin, and uptake foliar retention, (2006) Spray C. & Gauvrit M., B. Chauvel, Grangeot, Goeden, R.D. & Andres, L.A. (1999)Three rece This isarticle by protected copyright.reserved. Allrights Gross, K.L. (1990) A comparison ofmethods Guo, J.-Y., Zhou, Z.-S., Zheng, X.-W., Chen, H.-S., Zhou,X.-W.,(2011) Control J.-Y., Wan, Y.-H. &Luo, Z.-S., Zheng,Guo, F.-H. Grewling, Ł., Nowak, M., Jenerowicz, D., Szyma Hartmann, H. ragweed H. Hartmann, & Watson,to common( (1980)Damage A.K. Hanson, C.G. &Mason, J.L. (1985)Birdseed aliens in Britain, New Zealand New group. this of inwith Ambrosiinae North America in relation Handbook of biological control habitat models. DOI dx.doi.org/10.1016/j.tree.2014.02.009. shift niche studies: insights biological from invasions. potentials. weeds Assessingperiod of germination temperatures potential with base andbase water 11 common ragweedof ( behavior ofEcology Journal of translocation glufosinate and glyphosate in 13 EAACI-WAO EAACI-WAO World Allergy Congress 2013, Milan, Italy, pp. 686. ofconcentrations ragweed pollen and Amb a 1 in Poznarecorded Bogawski,P.,Ł., Šikoparija, (2013) Skjøth, B., &Smith, M.C. Atmospheric A. 152–162. efficiency of leaf beetle, artemisiifolia 1063. caused bycaused the white rust fungus ( , 217–223. , 933–944. Weed Research Weed (ed R.R. Scott), pp. 565–573. CSIRO, Melbourne, Australia. , at different, growing stages. 8th International onBiological 8th International Symposium Control ofWeeds, Canterbury, Ecology Letters , 78 , 1079–1093. Ophraella communa , 53 , 76–87. (ed TWFisher), pp. 884–885. Academic Press,San Diego. , Ambrosia artemisiifolia Ambrosia 8 Albugo tragopogi , 993–1009. Biocontrol Science andTechnology el, el, C., Munier-Jolain, N. & Colbach, N. (2013) , on the invasive common ragweed, ragweed, common invasive the on , the seed weight and burial depth on the seed onthe depth burial weight seed and the for for estimating seeds numbers in the soil. Ambrosia artemisiifolia Ambrosia ń nt successes outside of North America. In: to the biologicalto the control of weedymembers ies offering simple distribution: morethan ska, A., Czarnecka-Operacz, M., Kostecki, ned from studies of the insects the studiesof from associatedned ). Weed Science

). ). Trends inEcology andEvolution Weed Biology and Management Weed Biology Watsonia , Ambrosia artemisiifoliaAmbrosia ń 28 (Poland), 2010-2012. . , , 632–635. Weed Research 15 , 237–252. , 21 Ambrosia , 1049– , , 46 ) , , , Accepted Article This isarticle by protected copyright.reserved. Allrights a vegetation: dominance in apical of cost for Testing (1996) L.W. Aarssen, & D.L. Irwin, Igrc, J., F.V. DeLoach, and (1995) Zlof, Release C.J.establishment of & D.,Horváth, Kazinczi, & Keszthelyi,( G.S. répabarkó (2014)Akarcsú Hodgins, K.A. &Rieseberg, L.Geneti (2011) Bunce, (1999) & R.G.H. D.B. J.O., Roy, Mountford, M.O., Hill, I.(2014) surveyHeap herbicideresistantThe international weeds. of URL Jehlík, V., Dostálek, J. & Zaliberová, M. (2005) Spreading of adventive plants on river banksriver plantson adventive of (2005)Spreading &Zaliberová, M. Dostálek, V., J. Jehlík, Jehlík, V.(1998) Ciziexpanzivni plevele Ceske republiky a Praha. Slovenske republiky. Jacob,D., Petersen, J., Eggert,B., Alias, A., Christensen, O.,Bouwer, L., Braun,A.,Colette, Jenser, G., Kiss, B. &Takács, (2009) A. artemisiifoliaAmbrosia isa joint hostof tomato Joly, M.,P., Bertrand, Gbangou,R.Y., Lavoie, White,M.-C., Dubé,& J. (2011) Paving the C. nigrosaturatus Biology Evolutionary and native ( common ragweed introduced field study of three species. species. three of study field ( suturalis 374. Transport and the Regions, HMSO, London. 2. Annex Technical Vol. ECOFACT, plants British for http//:www.weedscience.com. Accessed 22February 2014. Haensler, Hempelmann, A., N.,Jones, C.,Keuler, K., Kovats, S., Kröner, N.,Kotlarski, S., A., Déqué, Georgievski, M., G., Georgopoul harbours. harbours. in the Elbe Czechthe RepublicandDanube River of outside Slovakiaof the in River research. ch high-resolution climate EURO-CORDEX: new (2013) P. Valentini, C., J.-F., R., B. R., Soussana, & Teichmann, Yiou, Vautard, Weber, S.,Rounsevell,D., Rechid, C.,Radermacher, Samuelsson, Somot, K., Radtke, P., M., Kriegsmann, A.,Martin, Meijgaard, E., E., Moseley, C.,Pfeifer, S., Preuschmann, S., occidentalis vectors, its and virus(TSWV) wilt spotted Magyarországon). közös vektorainak bronzfoltosság gazdanövényeés paradicsom (TSWV) vírus Ambrosia artemisiifolia Ambrosia F. (Coleoptera: Chrysomelidae) in of control Croatia for ragweedcommon Regional Environmental Change Thaiszia (Pergande) in Hungary (A parlagf , Goeze, 1777), a parlagf Goeze, 1777), a , , Növényvédelem 15 , , 35–42. 24 L.). L.). , 2731–2749. Annales BotaniciAnnales Fennici Biological Control, , 45 , 435–437. ű természetes ellensége. Ambrosia artemisiifolia Ambrosia , 14 , 563–578. ou, E., Gobiet, A., Menut, L., Nikulin, G., Nikulin, L., A., Menut, Gobiet, ou, E., Thrips tabci c differentiation of in life-history traits

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Accepted Article This isarticle by protected copyright.reserved. Allrights Reinhardt,F.,Herle, M., Bastiansen,Stre F.& Raynal, & D.J. Bazzaz, (1973)EstablishmF.A. Raynal, D.J. &Bazzaz, F.A. winter (1975)Interference of annuals with Pinke, G., Karácsony, P., Botta-Dukát, modelling biological niche B. (2014)UsingPetitpierre, to environmental understand of Payne,re-evaluation (1964) Agenusthe W.W. Pyšek, P. & Hulme, P. dynamics (2005)Spatio-temporal of plant-invasions: Linking pattern Mosyakin, (2006)Deliberateand unintentionalS.L. & Shevera, M.V. V.V., Protopopova, Prank, M., Chapman,D.S., Bullock, J.M., Belmonte, J., Berger,U., Dahl, A., Jäger, S., Poltavsky, A.N. & Artokhin, K.R. (2006) S.T. role Baskin, Pickett, (1973)The & J.M. Kueffer,Petitpierre, B., C., Broennimann, O., Randin, C., Daehler, C. & A.Guisan, (2012) Pyšek, Pyšek, P., Sádlo, J. & Mandák, B. (2002)Catalogue alien plantsof of the Czech Republic. Poppendieck, H.H. (2007) DieGattungen artemisiifolia Alien Species in Germany. J.W. Goethe University Frankfurt/Main, Frankfurt/Main, UniversityJ.W.in Goethe Germany. Species Alien soil. prairie and forest of Pest Science Pest of artemisiifolia ofbehavior invasions in a changing world.PhDthes Arboretum Arnold to process. 17–33. of Ukraine. alien study case the of weeds: of invasive flora a introduction 182-183 pollen ragweed release and dispersion in Europe. Severova, E.,I., Sikoparija, B. &Sofiev, model M. (2013) forecasting for Anoperational Kovtunenko, I., Magyar, D.,Niemelä, S., Rantio-Lehtimäki, A., Rodinkova, V., Sauliene, zuHamburg Vereins derAmbrosia-Bekämpfung. zur Hinweis Problematik einem the south of European Russia. invaders. plantamong terrestrial shifts are rare niche Climatic Preslia , , 43–53. 74 , 97–186. Ecoscience Ambrosia artemisiifolia Ambrosia in early successional fields. and other weeds to the management of Hungarian sunflower crops. management sunflower the Hungarian other of to weeds and , 86 , , 621–631. 65 , 23 , 401–438. , Ecology 12 , 53–70. , 302–315. Phegea , 54 , 1335–1341. . Bulletin of theBulletin BotanicalTorrey Club , 34 Tarachidia candefacta is, University of Lausanne, Lausanne Ambrosia Z. & Czúcz, B. (2013) Relating (2013) B. Czúcz, Z.& , 41–43. Ecology it, B. (2003) Economic Impact of the Spread ofEconomicof theSpread Impact it, B.(2003) ent of ent of successionalearly populations onplant of temperature and light in the germinationinthe andlight of temperature und , Ambrosia 56 Agricultural and Forest Meteorology , 35–49.

Iva (Compositae) inHamburg, mit (Compositae). (Compositae). Science (, ) in (Lepidoptera, Berichte des Botanischen , 335 , , 1344–1348. Euphytica 100 Journal of the of Journal , 165–170. Ambrosia Ambrosia Journal Journal , 148 , ,

Accepted Article Rybnicek, O., Novotna,O., Rybnicek,E. B.,Rybinickova, S.L.,Rousonelos, Lee, R.M., Moreira, M.S. Richter, R., Berger, U., Essl, S., Dullinger, (2013b) F. invasive G. &Vogl, Spread of Rich, T.C.G. (1994)Ragweeds ( Reznik, effects S.Y.(1991)The This isarticle by protected copyright.reserved. Allrights (2008)Basetemp G. I.&Pignata, Sartorato, Sang, W.,Liu, & X. Axmacher, and J.C. emergence (2011) Germination of Sheeley,S. & Squiers, E.R. P.E., Rothrock, Saint-Louis, S., DiTommaso, A. & Watson A.K. (2005) A common ragweed ( ragweed & WatsonA.K.A DiTommaso, common A. Saint-Louis, S., (2005) (2000) ed. J.S., Rodwell, Leitner, F., Essl, S., Dullinger, R., Richter, Reznik, S.Y., Spasskaya, I.A., Dolgovskaya, M. (Asteraceae) on populations of onpopulations of (Asteraceae) 737–743. artemisiifolia artemisiifolia Republic. ALS- and PPO-inhibiting herbicides. Characterization of a common ragweed ( Club Botanical of seedbank Ecology Applied ragweed: Climate change, management andhow toreduce allergy costs. insuitability weed management programs. Rector), pp. 614–619.International, CAB Wallingford. M.H. (eds Julien, R. Sforza, M.C. Bon, H.C. Evans, P.E. Hatcher, H.E. Hinz & B.G. Ambrosia artemisiifolia distribution, current abundance and implication for ofbiological control ragweed,common Oecologia http://www.nobanis.org/files/EconImpactNeobiota.pdf. Proceedings of the of 5 Proceedings 26 Vegetation of Open Habitats. The ragweed leafbeetle , 125–133. Aerobiologia , 88 Ambrosia artemisiifolia Ambrosia L. underchanging conditions in environmental China. linuron. to Québecresistant southwestern ) biotype in , 204–210. , 120 , 50 , 417–422. , 1422–1430. th , International Weed Science Congress (ed. International Weed Weed International (ed. Congress Science Weed International L. L. 16 British Plant Communities, Vol. Vol. Communities, Plant British Zygogramma suturalis , 287–290. 7th International Symposium7th International on Control Biological of Weeds Cambridge University Press, Cambridge, UK. Ambrosia of feedingof damageinragweed Zygogramma suturalis L. and Weed Science Weed L.) L.) in Britain. M. & Vogl, G. (2013a) How to account for habitatfor account (2013a) Howto G. Vogl, M. & & & Rybnicek, K. (2000) Ragweed Czech inthe Ambrosia artemisiifolia Ambrosia erature estimation of 21weed and species.crop Biological Invasions (1993) Heterogeneity and size of a persistent size a of (1993) Heterogeneity and Setaria faberii Y., Volkovitsh, & Zaitzev, M.G. V.F. (2007) , VanGessel, M.J. & Trane, P.J. (2012) P.J. & , M.J. VanGessel, Trane, F. (Coleoptera: Chrysomelidae) inRussia:F. , 60 Grana

, 335–344. (Coleoptera, (Coleoptera, Chrysomelidae). 5: Maritime Communities and Communities 5: Maritime Herrm. Herrm. , 33 , , 38–43. 15 ) population resistant to Ambrosia artemisiifolia , 657–669. Plant Species Biology Species Plant Weed Technology Weed Bulletin the Torrey of Journal of Journal Ambrosia Ambrosia Ambrosia , 19 , ,

Accepted Article Sheppard, A.W., Shaw,Sheppard, R. A.W., ( &Sforza, R.H. Schröder, G. & Meinlschmid, E. (2009) (1993) R.T. Koide, & R.P. Schreiner, Soluji This isarticle by protected copyright.reserved. Allrights weedannual of some fate the and changes (1974) Dormancy L.M. & Wax, E.W. Stoller, Simard, M.J. &Benoit, Distributionweed,anallergenic(2010) abundance and of D.L. Smith, M., Skjøth, A.,C. Myszkowska, D., Puc, M., Stach, A., Balwierz, Z., Chlopek, K., ragweed: ACommon (2013) B. Šikoparija, & G. A., C. Karrer, Skjøth, Cecchi, M., L., Smith, Singer,B.D.,Ziska, L.H., Frenz,D.A.,Gebhard Shrestha, A., Erivelton, S.R., Thomas, &Swanton, A.G. C. J. (1999)Modeling germination Shin, H.D. (2000)Erysiphaceae of Korea. Suwon, Šikoparija, B., Smith, Skjøth,M., C.A., Radiši Grewling, Radiši A., Dahl, Sommer,J., Ł., C.A., K., Kübler, B., Skjøth, Šikoparija, Alm atmospheric CO Canada. ( ragweed common Weed Research inEurope:a reviewcontrol of opportunities, Gesunde Pflanzen Ambrosie( Beifußblättriger Phytologist Science Society), p. 274. Vancouver, Canada. seeds inseeds the soil. to Poland. of transport & Brandt, I.J. Kasprzyk, (2008)Long-range K., Piotrowska, threat to environmental healthEurope.in Journal ofBiometeorologyJournal ThePannonian Plain as (2009) a source of of shoot-radicleand elongation Science Technology.and composition and biological activity of the acetone extract of extract acetone the activityof biological and composition ( ragweed common in content 1 Pannonian Plain. of transport long distance for M. (2013)Amechanism Smith, & ć , L.,Vukovi S., Sukdolak, , Canadian of Journal Plant Science Agricultural andForestMeteorology , 123 , 2 , 107–113. 46 concentration. Weed Science AgriculturalForest and Meteorology , , 93–117. 61 Ambrosia artemisiifolia , 135–150. , 53, Ambrosia artemisiifolia , 263–272. Ambrosia artemisiifolia Ambrosia 22 Functional PlantFunctional Biology ć Ambrosia artemisiifolia , N.,, Ni , 151–155. Mustards,mustard oils and mycorrhizas. Environment International 2006) Top 20 environmental classicalweeds for Top 20environmental 2006) ć , Ambrosia iforovi , , D.E. & J.G.Straka, (2005)Increasing Amb a 90 L.), southern inruralsettings Québec, of Untersuchungen zur Bekämpfung zur Untersuchungen von ć , P., Milkovska, S., Šimi , 549–557. regulations and other barriers to andotherregulations barriers to adoption. , Korea: National Instituteof Agricultural 148, ć , N. & Stani 1402–1411. polleninthe Balkans.

, L.) mit herbiziden Wirkstoffen. herbiziden Wirkstoffen. mit L.) . 180, Weed Science , ) pollen as a function of rising rising of function asa ) pollen 32, 112–117. 667–670. Ambrosia artemisiifolia Ambrosia ć , S. (2008) Chemical (2008) , S. , 61, , 47 115–126. ć , S. & Brandt, J. , 557–562. Ambrosia pollen from the from pollen International pollen pollen ć New New , P. L. L. Accepted Article Stinson, K.A. & Bazzaz, F.A.K.A.(2006) CO2 Stinson, &Bazzaz, en Steyermark, J.A. (1963) Skjøth,Smith, M., Stach, A., (2007) C.A. & Brandt, J. Examining This isarticle by protected copyright.reserved. Allrights Stojanovi Taylor, J.B. Loux, M.M., Harrison, S.K. & Clot, C., Lambelet, Ke B., P., Taramarcaz, Swanton, Shrestha,C.J., Knezevic, A., S.Z.,Roy,& R.C. B.R. Ball-Coelho, (2000)Influence Storkey, Stratonovitch, J., P., Chapman, Spangenberg, J.H.,Bondeau, A., Carter, T.R.,Fronzek,Jaeger, S., J., Jylhä, Kühn,I., K., Song, J.S. &Prots, of B. (1998) Invasion Stace, C. (2010) Stace, C. Teshler, M.P., DiTommaso, A., Gagnon, J.A. & Watson, A.K. (2002) Biometeorology Poznan usingback-trajectory (Poland) analysis. at competing stands stands competing of UK. Cambridge, pollen. pollen. 163. common ragweed ( common Weekly this healthinvasion? will its Switzerland resist risks: and progression SciencePlant tillage onvertical type seedbank distributionweed in asandyof soil. Europe. in plant allergenic invasive effect the predicting to approach process-based 31–36. candefacta Biogeography investigating risks for J. (2012)Scenarios tobiodiversity. Settele, & A.,Reginster, Paul, Rounsevell,I., I., Omann, Sciences ofBiological Journal Carpathians and Mts. Ukrainian the Tran L., commonL., ragweed (Asteraceae). B and herbicides. alternative Inhibiting ć , Journal oftheJournal Serbian ChemicalSociety , D.V., 135, 538–548. , , 80 21 Đ , New Flora of the British Isles oftheBritish Flora New 51, , 455–457. ur , 5–18. č Ambrosia artemisiifolia 275–286. i ć Flora ofMissouriFlora Ambrosia artemisiifolia , S.B., Orlovi (Hübner, 1831) (Lepidoptera, (Hübner, Noctuidae) u Srbiji. , 2 , 209–216. PLoS One ć , S., Kereši, T. & Gali & T. Kereši, S., , Weed Technology iological control programmesiological inCanada control . Iowa State University Press. Press. University State . Iowa imer, C. & Hauser, C. (2005) RagweedC. ( Hauser, & C. imer, D.S., F.Vidotto, &Semenov, M.A. (2014)A Ambrosia artemisiifolia Ambrosia Regnier E. (2002) Response ALS-ResistantRegnierof E. (2002) ) and giant ragweed ( ragweed ) andgiant scarpathian plain (Central Europe). scarpathian plain (Central Europe). richmentreduces dominance reproductive in , (common ragweed). M., Sykes, Schweiger,M.T. O.,Stocker, A., 9 of climate change on the distribution of an distribution of ofon the climate change ,

, e88156 73 , 3 , 1039–1049. rd Cambridgeedn. UniversityPress,

, 16 , 815–825. ć , Z. (2011), sovice Prvi nalaz International Journal of Ambrosia trifida Ambrosia Ambrosia L. (Compositae) in the(Compositae) in L. Ambrosia artemisiifoliaAmbrosia Oecologia Canadian Journalof Global Ecology andGlobal Ecology pollen pollen episodes Biljni lekar Swiss Medical , Ambrosia 147 ) to ALS- ) to (ed J.T. J.T. (ed Korean Korean , 155– , 39 ) ,

Accepted Article Toole, H.E. Brown, resu (1946)Final E. & Tokarska-Gudzik, B., Bzdega, K., Koszela, K., Zabinska,Krzus, Sendek, M. I., Sajan, & B., This isarticle by protected copyright.reserved. Allrights Thompson, K., Bakker, J.P. & Bekker, R.M. (1997) Týr, Š., Vereš, T. & Lacko-Bartošová, M. (2009) Occurrence of common ragweed ( Tosi, A., B.,Wüthrich, M. Bonini, Piet & Vogl, G., Smolik, M., Stadler, L.-M., Leitner, M., Essl, F., Dullinger, S., Kleinbauer, I. & Kleinbauer, Dullinger, S., F., Essl, Leitner, M., L.-M., M., Stadler, Smolik, G., Vogl, Tropicos (2014) Missouri Botanical Garden(2014) Missouri Botanical Tropicos Vanky, K., S., Toth, Gonczol,J. &Revay,A. (1988) Furtherspecies of six Ustilaginales,new of L.,Bohár,Vajna, G. &Kiss,(2000) Firstreport L. inHungary L. by ragweed on (2002)DownyVajna, epidemic caused common mildew Vitalos, M. & Karrer, G. (2009) Dispersal of Dispersal G. (2009) Karrer, & M. Vitalos, Vidotto, F.,Vidotto, F. Tesio, (2013)Allelopathic & A.F. Vidotto, of effects Varga, P., Beres, I.&Reisinge &Kovacs Kazinczi, Varga,G., P., I. Beres, Vincent, G. & Cappadiocia, M., (1988) Charac (1988) Cappadiocia, M., G. & Vincent, selected aspectsof biology. of Agricultural Research of invasiveplant Allergenic (2011) A. methodology, densityand longevity Huber), pp. 290–294.Publishing, CABI New York. contribution ofcontribution traffic and mowing machines. artemisiifolia Accessed 26 February 2014. Italy.northern Ambrosia causing epidemics on ragweed.common 576. Hungary.to Plasmopara halstedii ragweed, ragweed, inL. the invasive process. on the yields of maize in different years. 701–704. artemisiifolia Ambrosia sensitisation and sensitisation Ambrosia artemisiifolia Acta Botanica Hungarica L.) in field crops in the Slovak Republic. Swiss Medical Weekly Medical Swiss . Plant Pathhology in additive experiments. experiments. in additive , 72 Crop Protection Biodiversityand Research Conservation , 201–210. Ambrosia r, P. (2002) The r, P.competitive effect of weeds three dangerous , and giant ragweed, ragweed, giant and , . Cambridge University Press, Cambridge ,

allergy: 20-yearA study 141 34 Ambrosia artemisiifolia , Növényvédelem lts of buried Durvelthe seed experiment. Plant DiseasePlant , 193–208. 51 , w13253. , I. (2006) Competition between sunflower sunflower and between , I. (2006) Competition . URLhttp://www.tropicos.org/Name/2701648. , ragalla-Köhler, B. (2011) Time lag between lag(2011) Time B. ragalla-Köhler, , 809. , 809. 54 Neobiota terization of terization hybrids reciprocal common of Ambrosia artemisiifolia , 161–167. The soil seed banks seed ofThe soil northwest Europe: Cereal Research Communications Research Cereal

Phyllachora ambrosiae , , Herbologia A. trifida A. 84 8 , , 53–60. 38 , 489. , 219–226. L. in Poland: threat and . (1989-2008) in Legnano, Legnano, in (1989-2008) Weed Science, Weed , Ambrosia artemisiifolia 10 , 21 , 1–9. seeds alongroads: , 39–48. in Europe Europe in

Ambrosia 36 Journal Journal , 574– , , 34 ,

Accepted Article Webb, C.J. (1987) Checklist of dicotyledons na (1987)Checklist Webb, C.J. ofWeaver, lamb's-quarter (2001)Impact S.E. This isarticle by protected copyright.reserved. Allrights Webster, T.M., Cardina, J. &White, A.D. (2003) seed rain, seedbanks, Weed soil and Webster, T.M. &Nichols, R.L. (2012) Changes White, & J.F. D.I. Bernstein, (2003)Key pollen allergens inNorth America. Wayne, of Foster,allergenicS.,Production P., & J., F. Epstein, P. (2002) Connolly, Bazzaz, Wang, D. & Zhu, X. (1996) Research onallelopathy of H. &Voglmayr, A. ( Riethmüller, Watanabe, O., Kurokawa, S., Sasaki, H., Nishida, T., Onoue, T. & Yoshimura, Y. (2002) F.H., MaWan, Gui, J., &You,L. J.Y. Wagner, &Beals, T.F. ( (1958) W.H. ragweeds Perennial Lippe, (2013)Human- von M., J.M., &Wichmann, M. Bullock, I., der Knopp, Kowarik, T. Willemsen, R.W. (1975) Effect ofWillemsen, stratification R.W. (1975)Effect temperature onand germination temperature seedling recruitmentseedling in no-tillage crop rotations. Science to 2008/2009. the United crops 1994/1995 agronomic southern States: of major diffusion. habitat,and change climate effects of ragweed: of Modelling spread the (2008) J. Peterseil, (Compositae) subfamily(Compositae) Asteroideae. maize of and soyabean in Ontario. ofAllergy,Annals Asthmaand Immunology Science Ecologica Sinica 473–478. against ragweed, new, ofadescription taxon. intermediate Allergy AsthmaandAllergy Immunology by ( pollen ragweed Geographic scale distribution and occu strenuana mediated dispersal of seeds by the airflow of vehicles. 75–85. rusts)blister based on LSU germination and the induction of secondary of induction seeds. andthe dormancy in ragweed common germination , , 60 48 (Lepidoptera: Tortricidae) and and Tortricidae) (Lepidoptera: European Journal ofPhysics , 145–157. , 440–450. , 16 Ambrosia artemisiifolia Ambrosia artemisiifolia Ambrosia , 11–19. rDNA andsequence oospore data. 2006) Phylogenetic relationships of , 91, Canadian Journal ofPlantScience 425–435. New Zealand Journalof Botany , S. (2003) Integrated control effects of of effects S. (2003)Integrated control 161 Rhodora rrenceof invasive pattern weeds. (Compositae). , s, ragweeds, greencommon onyieldand foxtail L.) is increased in CO , 167–173. 8 Ostrinia orientalis Ostrinia , 279–282. turalised NewZealand.in Asteraceae 18. in theprevalence of weed species in the Weed Science Weed , 60 , 177-204.

PLoS One Acta EntomologicaSinica Ambrosia Ambrosia artemisiifolia Ambrosia , Mycological Research Mycological 51 (Lepidoptera: (Lepidoptera: Pyralidae) , 8 , 569–575. 2 , e52733. -enriched atmospheres. -enriched atmospheres. Albugo , , ) in Michigan, ) inMichigan, with 25 81 , 489–501. , 821–828. species (white Grassland Annals of Epiblema Epiblema . , Weed , Acta Acta 110 46 , , Accepted Article Ziska, L.H., Gebhard, D.E., Frenz, D.A., Faulkner, S., Singer, B.D. & Straka, J.G. (2003) Filley, W., Elder, M.A., N., Tierney, Dalan, D., C., Rogers, Knowlton, K., L.H., Ziska, survival,on of temperature Effects (2010) J.Y., Wan, F.H. Guo, H.S.Zhou, & Chen, Z.S., Wopfner, Gadermaier, N., Egger, G., Asero,M., Appendix S1. Appendix be Supporting informationAdditional may intheonlinefound of this version article: information Supporting This isarticle by protected copyright.reserved. Allrights Wodehouse, R. P.(1971) Willemsen, R.W. & Rice, E.L. (1972) Figure S5. Figure S4. Figure S3. Figure S2. Europe. ininvaded S1. Figure climate, as predicted by SDMs. artemisiifolia American Journal ofBotany of Allergyof andImmunology of allergens Thespectrum andmugwort inragweed (2005) Pollen. Journal ofAllergyImmunologyJournal and Clinical as harbingersCities ofclimate change: co 4251. America. North central warmingRecent associated by latitude withincreased length ofragweed season pollen in J., (2011) Portnoy, F., Coates, D. Vrtis,Frenz, J.A., Patz, Bielory, G., R.F., L.& Fulford, J.,Shropshire, Ford,L. B., Hedberg, C., Fleetwood,P., Hovanky, K.T., Kavanaugh, T., (Asterales: Asteraceae). potential aChrysomelidae), agent biological control against longevity, and fecundity of development, The potential binarized distribution of SDMs. inofThe usedrelativeas predictors variables importance of The spread Occurrence of Appearance of Modelling the spread of . American Journal of Botany of American Journal Hayfever Plants Hayfever Ambrosia artemisiifolia Ophraella communa Physiological Ecology Proceedings of the National Academy of Sciences of theNational Academy Proceedings of Ambrosia artemisiifolia Ambrosia , , 138 62 , 1–5. , 337–346. Ambrosia artemisiifolia. , 2 Mechanism of seed dormancy in nd mmon ragweed, urbanization, and public health.mmon ragweed, urbanization, andpublic , ed. Hafner Publishing,New York. in Europe in 2013. in in Europe 59 , 111 R., R., C., Ebner, Jahn-Schmid, B. &Ferreira, F. in andEasternCentral Europe. , 248–257. , A. artemisiifolia A. 39 , 290–295. Ophraella communa , 1021–1027. populations in habitats that aremost in populations

under andcurrent future Ambrosia artemisiifolia International Archives (Coleoptera: , 108 Ambrosia , 4248–

Accepted Article This isarticle by protected copyright.reserved. Allrights Table 1. Tables the phytosociological four classesinwhich it is typical northern Italy, and eastern France) (Nitzsche 2010 Romania, Germany, relevésphytosociological from 220 on (based Eastern Europe and Taraxacum Taraxacum viridis Setaria pumila Setaria aviculare Polygonum major Plantago lanceolata Plantago Artemisietea lupulinaMedicago perenne Lolium Stellarietea Lactuca serriola repens Elymus crus-galli Echinochloa Daucus carota canadensis Conyza arvensis Convolvulus album Chenopodium vulgaris Artemisia millefolium Achillea Species Tripleurospermum inodorum inodorum Tripleurospermum repens Trifolium pratense Trifolium The plant species Theplant mostco sect . Ruderalia agg agg . . mmonly associated with x x x x x x x x , modified). Representationis indicated, for

Ambrosia artemisiifolia Ambrosia x x x x Arrhenatheretea Molinio- x x x x x x in Central in Central Polygono-Poetea Plantaginetea, x x Accepted Article Index Fungorum (http://www.indexfungorum.org/Index.htm) (http://www.indexfungorum.org/Index.htm) Index Fungorum range (modified inEurope Gerber from This isarticle by protected copyright.reserved. Allrights Table 2. I Taxon Country Source ecpde Adelphocorislineoletus Coreusmarginatus spumarius Philaenus Cercopidae C hyoeia Mordellidae Coniocleonusnigrosuturatus Tanymecuspallidus Sitona suturalis Phyllobius pyri Lixus Curculionidae Zygogrammasuturalis Ophraella communa Galeruca tanaceti sericus Cryptocephalus Chrysomelidae dahli Agapanthia Cerambycidae H Morellistena NSECTA OLEOPTERA OLEOPTERA ETEROPTERA ETEROPTERA s. fre uolva 16 formerYugoslavia sp. Herbivores and pathogens recorded on and Herbivores recorded pathogens

sp. Russia 16 16 Russia sp.

L fre uolva 16 Yugoslavia former L. Steph. former Yugoslavia 16 16 Yugoslavia former Steph. Rctr ugr 12 Hungary Richter L. former Yugoslavia, Yugoslavia, former L. L. Russia, former Yugoslavia, Yugoslavia, former Russia, L. R Rsi,fre uolva 16 Russia, formerYugoslavia R. L Hnay former Hungary, L. Le Sage Le F. L fre uolva 16 Yugoslavia former L. Gee ugr 13 Hungary Goeze

Goeze

et al. 2011). Fungal taxonomy is in accordance to Fungalis taxonomy2011). Slovakia Slovakia Yugoslavia, Sweden Hungary Hungary 22 10, 17 Russia, Croatia Switzerland Italy, Slovakia 9

Ambrosia artemisiifolia

in itsintroduced 16, 27 13, 28 16, 27 Accepted Article This isarticle by protected copyright.reserved. Allrights L sp. Psyllid Psyllidae Parthenolicaneum corni Diaspididae Membracidae Eupteryx atropunctata molliculaEmelyanoviana Cicadella viridis Cicadellidae Protaphis rmia Ostrinianubialis Crambidae H Eurydema ornatum Eurydema oleraceum interrupta Eurydema oleraceum flanata pratensisLygus rugulipennis Lygus Myzus persicae Brachycaudushelichrysi Aphis Aphis fabae Aphididae (Kaltenbach) Stictocephala EPIDOPTERA EPIDOPTERA OMOPTERA OMOPTERA s. fre uolva 16 former Yugoslavia sp.

sp. Russia 16 16 Russia sp. Sweden 28

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(oz) ugr 13 Hungary (Goeze)

Buh Rsi 28 Russia Bouché (oea) ugr 13 Hungary (Boheman) Sh. omrYgsai 16 Yugoslavia former Schr. R. omrYgsai 16 formerYugoslavia Ry.

Slovakia 27 Sweden 2 Hungary Slovakia 27 Slovakia 27 Yugoslavia Yugoslavia Hungary, former Yugoslavia, Slovakia

13, 27 2

Accepted Article This isarticle by protected copyright.reserved. Allrights rlia T Leptophytesbosci Tetrixundulata Tettigidae Oecanthus pellucens Gryllidae Pholidoptera Decticinae Chortippus Chortippusparalellus Acrididae O 16 Tarachidiacandefacta 16 Scotia ipsilon Peridromasaucia Hypena proboscidalis Helicoverpa armigera former Yugoslavia Chloridea scutosa former Yugoslavia Autographa gamma Autographaconfusa Noctuidae Orgyarecens Lymantriidae Unidentified Unidentified Cosymbia Geometridae hiia Aeolothrips intermedius Thripidae HYSANOPTERA RTHOPTERA RTHOPTERA

s omrYgsai 16 formerYugoslavia sp

s. fre uolva 16 formerYugoslavia sp.

s. fre uolva 16 former Yugoslavia sp. Hn fre uolva 16 former Yugoslavia Hbn. Rt. usa 14 Russia Rott.

Sr. omrYgsai 16 Yugoslavia former Serv. Fe. omrYgsai 16 formerYugoslavia Fieb. Hn fre uolva 16 Yugoslavia former Hbn. Shf. usa 16 Russia Schiff. L Rsi,fre uolva 14 Yugoslavia former Russia, L. Seh Rsi 14 Russia Steph. So. omrYgsai 16 former Yugoslavia Scop. Zt. omrYgsai 16 Yugoslavia former Zett. L fre uolva 16 Yugoslavia former L. (Hübner) Hübner Bagnall

tl,Soai 27 Italy, Slovakia usa eba 20, 23 Russia, Serbia Hungary 11 Hungary

Accepted Article This isarticle by protected copyright.reserved. Allrights & De Toni halstediiPlasmophora angustiterminalisPlasmopara O O B A Thrips tabaci Thrips nigropilosus fuscipennis Thrips Crawford) Microcephalothrips abdominalis Haplothripsaculeatus occidentalisFranliniella Frankliniella intonsa D P Gray) S.F. Albuginaceae A A erncia F sp. Aceria Eriophyidae Tetranychus urticae Tetranychidae oroparaee Botryosphaeriaceae Peronosporaceae Thienes Pustula tragopogonis ERONOSPORALES UNGI OTRYOSPHAERIALES SCOMYCOTA LBUGINALES OMYCETES OMYCOTA CARINA OTHIDEOMYCETES

( Albugo tragopogonis

Lnea Hnay 11 Hungary Lindeman

aia ugr 11 Haliday Hungary Kc. usa 16 Russia Koch. zlHnay 11 Uzel Hungary

(Pers.) (Pers.) Tyo)Hnay 11 Hungary (Trybom) (Farl.) Berl. Berl. (Farl.) Fbiis ugr 11 (Fabricius) Hungary Prad)Hnay 11 (Pergande) Hungary Nvt Urie 8 Ukraine Novot. (D.C.) (D. L.

Hungary 11 Hungary ugr,Asra 6, 26 Hungary, Austria Hungary 24 Hungary

Serbia 29

Accepted Article This isarticle by protected copyright.reserved. Allrights H I avenaceum Fusarium Netriaceae H C Macrophominaphaseolina L Phoma Incertae sedis alternate Alternaria Pleosporaceae P Septoria epambrosia ambrosiaeSeptoria Mycosphaerellaceae S Leveillulataurica latispora ( latisporus Golovinomyces cichoracearum Erysiphaceae Plectosphaerellaceae E sclerotorium Sclerotinia cinerae Botrytis Sclerotiniaceae ( NCERTAE SEDIS NCERTAE Heluta( Golovinomyces cichoracearumvar. LEOSPORALES ORDARIOMYCETES EOTIOMYCETES RYSIPHALES APNODIALES YPOCREALES ELOTIALES Alternaria tenuis Erysiphe cichoracearumvar. s. ugr 3 Hungary sp. Erysiphe cichoracearum U. Braun) (U. Braun)U. Braun

es ugr 6 Pers. Hungary Nees) Nees) (é. .Anu fre SR 1 USSR former Arnaud (Lév.) G.

Hmi&N at Jpn 18 Japan Hemmi& N. Naito

(Fr.) Keissl D.F. Farr D.F. F..Sc. hn 15 China Sacc. (Fr.). (Lib.) de Bary Tsi od ugr 4 Hungary (Tassi) Goid

(DC.) V.P. (DC.)

DC.)

China 15 Hungary 5 Hungary Germany 7 7 Germany

Hungary 4 Hungary Mauritius Korea,

19, 22

Accepted Article This isarticle by protected copyright.reserved. Allrights (unpublished data); (28) Stephan Vajna (2002);(25)Vanky Artokhin (2006);(21) (1991); Reznik Müller-Schärer (2008); (15)Li(14) Kovalev (1971b); &Li Kiss Jenser, (11) & Zlof(1995); & DeLoach Igrc, Kazinczi(2014); (10) &Keszthelyi (9)Horváth, (2007); &Hayova Dudka Schwarzinger (1999);(6) (1999); (5)Bohár& (2) (3) BaskyBohár Sources: (1) Amano(1986); (2009); (unpublished data). E C P Verticillium dahliae E Donk Thanatephoruscucumeris Ceratobasidiaceae I Entylomapolysporum Entylomataceae A Sacc. Curtis) ambrosiae Phyllachora Phyllachoraceae Athelia rolfsii rolfsii Athelia ( NCERTAE SEDIS NCERTAE HYLLACHORALES NTYLOMATALES XOBASIDIOMYCETES ANTHARELLALES GARICOMYCETES Sclerotium rolfsi

( Rhizoctonia solani solani Rhizoctonia et al. (Curzi) Tu & (Curzi) C.C. Kimbr.

(2014); Orie (18)Naito (1940); (19) Sacc.)

Kleb. Kleb. Pc)Fr. Hnay 25 Hungary Farl.(Peck) (Berk. &M.A. (Berk. et al. J.G. Kühn) (A.B. Frank) (1988);(26)Voglmayr Riet & Takács (2009); (12) Kiss (2009), (2009); (12) Takács et al. (22) Shin (2000); (23) Stojanovi (23) (2000); Shin (22) (unpublished data); (29) Petanovi (29) (unpublished data); Bohár & Vajna (1996); (7) Braun (1995); (8) Vajna (1996);(7)Braun(1995); & Bohár (1993); (16) Maceljski & Igrc (1989), (17) Igrc (1989), & Maceljski (16) (1993); Hungary 6 Hungary Hungary 6 Hungary

Hungary 24 Hungary China 15

ux & Felix (1968); (20) Poltavsky & Poltavsky& (20) (1968); Felix ux & et al. et hmüller (2006);hmüller (27) P.Toth (13) Kiss, Rédei & Koczor &Koczor Rédei Kiss, (13) Kiss & Bohár (4) (2009); ć

et al. et ć (2011);(24) &Vidovi ć

Accepted Article = increasing; orange = stable; yellow = decreasing) =decreasing) yellow stable; = orange increasing; = todispersal,trendsthe andrelativetemporal contribution importance (red putative putative in range < (short 1 km;medium distance distance 1–100 km;longdistance >100 theirkm), This isarticle by protected copyright.reserved. Allrights Table 3. Pathway Spatial Spatial Pathway range Relative Contaminated soil (traffic) Vehicles Mowing machines machinery Agricultural seed fodder/birdAnimal seedsplanting for oilseeds,Grain, Human-mediated seed dispersal Hydrochory Barochory Natural seed dispersal Dispersal pathways Dispersal for

distance long medium / Short / Short distance distance Medium distance long medium / Short / distance Long distance /long Medium distance medium Short / Short distance

Ambrosia artemisiifolia importance importance Low Low High Medium High High EFSA Medium Medium Low (2010)

importance in trend Temporal Bassett & Song & Prots, in Europe. Shown are their spatial their Shown are inEurope. References Selected (1975) (1975) Crompton al. Taramarcaz (2009) Karrer & Vitalos (2009) Karrer & Vitalos (2011) Karrer al. Karrer (1998), (2007) Fumanal (2005) (2011) et al. et al. et et et et

Accepted Article This isarticle by protected copyright.reserved. Allrights (2013a,b) al. Richter (2008), Vogl al. Storkey (2014) et al. Chapman (2012) et al. Petitpierre on based MS; This al Cunze Scale Model Ambrosia artemisiifolia Table 4. . (2013) (2014) et al. et et

et et Summary ofmodels predicting effect the of climate change onthefuture spread of

Europe Europe km) (5x5 Europe km) (50x50 Worldwide (5’x5’) Europe (resolution) (3’x5’) (3’x5’) Europe Central apart) >100 km (points simulate spread to kernel dispersal niche with species coupling ‘Hybrid model’ rate. growth population positive with region as range Predicts dynamics. seedbank and reproduction nutrients, and water light, competition for representing simulator growth Mechanistic plant frost. lifecycle before completes ragweed where region as Range predicted photoperiod. and temperature by driven model phenology Mechanistic occurrences. Australia) Eurasia, (North America, and non-native native to fitted model distribution species Correlative occurrences. native to fitted model distribution species Correlative Approach Climate Other variables in Europe coldest annual the of temperature minimum seasonality, temperature temperature, Annual mean quarter. warmest of precipitation wettest quarter, of precipitation seasonality, precipitation quarter, warmest of temperature seasonality, mean temperature range, temperature diurnal Mean precipitation. precipitation. annual mean temperature, Mean annual precipitation. and Daily temperature temperature. maximum and Daily minimum seasonality moisture and moisture seasonality, annual precipitation wettest quarter, the of precipitation quarter, hottest annual the of temperature quarter, maximum variables variables – 2050, 2050, – 2080 IPCC roads. of length cover, land urban and Crop 2070 IPCC IPCC 2050 Photoperiod.

year End 2100 2100 2050 IPCC IPCC 2050 scenarios climate Future A1b A1b IPCC B2 A2, A1, range: upper (lower & A1b A1b A2a summary prediction Spread Potential variable. important most is the by far temperature Annual mean Ukraine). and Russia (e.g. southern range the of in drierparts populations current Extinction of Scandinavia. central and Britain southern as spread as far Northwards occurrences current multiple infilling from by Spread margin. in southern Little change Scandinavia. southern and Britain as spread as far northwards allow frosts delayed and summers Warmer range margin. southern of representation No Scandinavia. central as spread as far northwards allow modest frosts delayed and summers Warmer Siberia. and Europe northeastern in particular in change, in climate under globally increase will distrinbution Accepted Article This isarticle by protected copyright.reserved. Allrights al. Bullock (2012) (2012) et (5x5 km) km) (5x5 Europe in study region. region. in study to data on spread Fitted dynamics. distribution. distribution. current Calibrated against dynamics. simulate spread through trade to introduction ragweed and model extinction kernel, dispersal niche with species coupling ‘Hybrid model’ seasonality. temperature precipitation, and temperature season Growing rates. rates. import seed national cover, land urban and Crop

2080 IPCC IPCC 2080 +2.4 °C) +1.5 °C, A2a, B2A2a, A1b, change. climate under worse (upper range) ~50% and (lower range) is ~25% range promotes Europe Eastern in cropland Loss of for ragweed. dry and hot becomes too Europe Southern Scandinavia. southern and Britain as spread as far northwards allow summers Warmer spread. restricts and contraction Accepted Article This isarticle by protected copyright.reserved. Allrights Figures least one least the record ina10-kmsquareof Grid.one National ( 1. Fig. of the British Isles, usingDr A. Morton’s DMAP software. SocietyBotanical the of bycollected members records from mainly Ecology andHydrology, pre-1970.Mappednon-native by Harrower, Biological Records Colin Centre Centre, for

The distribution of of The distribution Ambrosia artemisiifolia Ambrosia in the British Isles. British inthe

+ ) non-native 1970onwards;( ) non-native

Each dot represents at dotrepresents Each

x ) Accepted Article areas with high level of allergies recorded (based on Déchamp, Méon & Reznik 2009). additional records (D.S. Chapman, unpublished da artemisiifolia Ambrosia 2. Fig. This isarticle by protected copyright.reserved. Allrights The distribution (colonized grid cells of 50 x 50 km size are shown in grey) of of in grey) 50 kmsizeare shown of 50x griddistributioncells The (colonized inEurope.Distribution data are based onBullock ta). Black points indicate heavily indicate infestedta). Blackpoints

et al.et (2012), with

Accepted Article modified). This isarticle by protected copyright.reserved. Allrights Fig. 3. Fig. 3’, c. 35km (5xMappingAustria’ ‘Floristic of project of the line) of (dashed grid cells number colonised Austria. Results are shown ascumulative the number of records (solid line) as and cumulative Time course of invasionTime course by 2 ). The two lines coincide (c) for fields (Essl, Dullinger & (2009,Kleinbauer Ambrosia artemisiifolia

in six different habitats (a-f) in (a-f) habitats different in six

Accepted Article (e) male racemes.© Essl (4x), B. Fumanal (1x), M. Smith (1x). seedlings ofsunflower ( 4. Fig. This isarticle by protected copyright.reserved. Allrights Appearnace of Ambrosia artemisiifolia Helianthus annuus Helianthus ), (c) juvenile plant, (d) female synflorescence and synflorescence (d) female plant, ),juvenile (c) (a) whole plant, (b) seedlings, together with together (b) seedlings, plant, whole (a)

Accepted Article This isarticle by protected copyright.reserved. Allrights and(c)pollen.Drawings by KrisztinaBíró©. 5.Fig. Male capitula of Ambrosia artemisiifolia: Ambrosia (a) synflorescence; (b) single male flower; male (b)single synflorescence; (a)

Accepted Article This isarticle by protected copyright.reserved. Allrights longitudinal section;4,achene; 5, syconium cross-section.Drawings byKrisztina Bíró©. seeded syconium viewed side; synconium the from 2, viewed from above; 3, synconium in 6. Fig. Female capitula capitula Female of Ambrosia artemisiifolia : (a) racemes (a)with achenes;: (b)1, one-

Accepted Article Fig. 7. Fig. This isarticle by protected copyright.reserved. Allrights ©. Bíró Krisztina Germination and seedling development of Ambrosia artemisiifolia,

Drawings by Accepted Article Fig. 8. Fig. This isarticle by protected copyright.reserved. Allrights artemisiifolia (c) Pupa; (d) (e)Content of Adult; sweepa net 10sweeps after in a field infested with

Ophraella communa near Milano (Corbetta, 24 September 2013). on Ambrosia artemisiifolia Ambrosia in Northern Italy: (a) Eggs; (b) Larva; Larva; (b) Eggs; (a) Italy: Northern in

A. A.

Accepted Article This isarticle by protected copyright.reserved. Allrights climate where predictions are notreliable. For details,see Appendix S1. using CSIROthe MK3.0 Global Circulation M (the scenario change A1b-SR climate severe and (c Theprojection d) climates. habitatwarmer and future suitabilityof is a shown for used to were (a) pr SDMs calibrate Australia non-native inEurasia and the occurrences distribution species and for in America North 9. Fig. World-wide potential distribution distribution potential of World-wide ES-scenario) for the year 2050 (c) and 2100 (d) 2050 theyear (c)and for ES-scenario) oviding potential distribution under current (b) undercurrent distribution oviding potential Ambrosia artemisiifolia odel. Shaded area represents area Shaded non-analog odel.

. Information about . Information Accepted Article In Eastern Europe; there are gaps there Europe; resulting are In Eastern pollen lack from of stationsa (Smith data Aeroallergen thestored in Netw European zones 200 andpresence/absence of informakm index of with flowering potential theyearsfor 2000-09.Themapisbased onthe annual mean pollen 10. Fig. This isarticle by protected copyright.reserved. Allrights modified). A spatial assessment of densitythe of naturalised Ambrosia artemisiifolia from 368 stations in Europe, simple interpolation, buffer ork database (https://ean.polleninfo.eu/Ean/). tion in Flora Europea. The map isbased on The map tion Europea. in Flora

Ambrosia artemisiifolia artemisiifolia Ambrosia populations et al. 2013, 2013,